Fossil Fruits and Seeds of the Middle Eocene Messel ...

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Fossil Fruits and Seeds of the Middle Eocene Messel biota, Germany Margaret E. Collinson, Steven R. Manchester & Volker Wilde

Content Introduction.......................................................................................................................... 2 Geologic setting and age...................................................................................................... 2 Modes of preservation.......................................................................................................... 3 Material and methods........................................................................................................... 4 Floristic composition........................................................................................................... 6 Comparison with diversity known from leaves and pollen................................................. 7 Fruit and seed biology, dispersal and animal diets.............................................................. 7

Soft to leathery fleshy fruit tissues........................................................................................................7 Vertebrate gut contents containing fruits and seeds...............................................................................8 Insect dispersal and feeding...................................................................................................................9 Dry fruits and seeds: dehiscence mechanisms and vertebrate diet........................................................9 Dry fruits and seeds as a vertebrate food resource: fibrous, tough, woody and resinous tissues..........9 Endosperm food reserves.......................................................................................................................10 Epizoochorous dispersal........................................................................................................................10 Wind dispersal.......................................................................................................................................10 Other dispersal mechanisms..................................................................................................................11 Dormancy and germination...................................................................................................................11

Taphonomic Considerations................................................................................................. 11 Growth habits and vegetation reconstruction...................................................................... 12 Climatic interpretations........................................................................................................ 13 Biogeographic considerations.............................................................................................. 13 Comparison with other fossil floras.......................................................................................................13 Comparison with modern day floras......................................................................................................16

Future directions for Messel palaeobotanical research........................................................ 16 Systematics.......................................................................................................................... 17 Family Doliostrobaceae Kvaček. ..........................................................................................................17 Family Alangiaceae DC.........................................................................................................................17 Family Altingiaceae Lindl.....................................................................................................................17 Family Anacardiaceae R. Br.................................................................................................................18 Family Apocynaceae Juss......................................................................................................................20 Family Arecaceae Bercht. & J. Presl. .................................................................................................20 Family Bignoniaceae Juss.....................................................................................................................21 Family Burseraceae Kunth 1824...........................................................................................................21 Family Cannabaceae Martinov. ...........................................................................................................22 Family Cyclanthaceae Poit. ex A. Rich.................................................................................................22 Family Cyperaceae Juss........................................................................................................................23 Family Elaeocarpaceae Juss..................................................................................................................23 Family Euphorbiaceae Juss...................................................................................................................24 Family Hamamelidaceae R. Br.............................................................................................................25

Family Icacinaceae Miers.....................................................................................................................27 Family Juglandaceae DC.......................................................................................................................32 Family Lauraceae Juss...........................................................................................................................33 Family Leguminosae Juss......................................................................................................................34 Family Lythraceae J. St.-Hil.................................................................................................................35 Family Magnoliaceae Juss.....................................................................................................................36 Family Mastixiaceae Calest.................................................................................................................36 Family Menispermaceae Juss................................................................................................................38 Family Myristicaceae R. Br..................................................................................................................46 Order ?Nymphaeales.............................................................................................................................46 Family Nyssaceae Juss. ex Dumort.......................................................................................................47 Family Pentaphylacaceae Engl.............................................................................................................47 Family Rhamnaceae Juss.......................................................................................................................47 Family Rutaceae Juss............................................................................................................................48 Family Sabiaceae Blume.......................................................................................................................49 Family Sapotaceae Juss.........................................................................................................................49 Family Simaroubaceae DC....................................................................................................................50 Family Tapisciaceae Takht...................................................................................................................50 Family Theaceae Mirb...........................................................................................................................50 Family Toricelliaceae Hu. .....................................................................................................................51 Family Ulmaceae Mirb..........................................................................................................................52 Family Vitaceae Juss.............................................................................................................................52

Incertae Sedis....................................................................................................................... 56 Acknowledgements.............................................................................................................. 76 References............................................................................................................................ 77 Tables................................................................................................................................... 86 Plates.................................................................................................................................... 95 Index.................................................................................................................................... 248

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Fossil Fruits and Seeds of the Middle Eocene Messel biota, Germany Margaret E. Collinson, Steven R. Manchester & Volker Wilde

Abstract A survey of the extensive fruit and seed collections from the Middle Eocene oil shale of the Messel Formation, at Messel Pit Fossil Site, a UNESCO World Heritage Site at Messel near Darmstadt, Germany, reveals at least 140 genera, representing more than 36 families. The flora includes occasional conifer remains (Doliostrobus scales) and numerous angiosperm remains. The following angiosperm families are represented (of which ten denoted “*” are new records for Messel): Arecaceae, Alangiaceae (*), Altingiaceae (*), Anacardiaceae (4 genera), ?Apocynaceae, Bignoniaceae, Burseraceae (*) (2 genera), Cannabaceae (*), Cyclanthaceae, Cyperaceae, Elaeocarpaceae (*), Euphorbiaceae, Hamamelidaceae (2 genera), Icacinaceae (6 genera), Juglandaceae (3 genera), Lauraceae (c. 4 morphotypes), Leguminosae (c. 5 morphotypes), Magnoliaceae, Mastixiaceae (5 morphotypes), Menispermaceae (17 morphotypes), Myristicaceae (*), ?Nymphaeales, Nyssaceae, Pentaphylacaceae, Rhamnaceae (*), Rutaceae (5 morphotypes), Sabiaceae (*), Salicaceae, Sapotaceae, Simaroubaceae, Tapisciaceae (*), Theaceae, Toricelliaceae (*), Ulmaceae, Vitaceae (7 morphotypes), plus 65 morphotypes of unknown familial affinity. The genera Berchemia, Mytilaria and Pleiogynium are here recorded for the first time from the Paleogene. The assemblage indicates a wide range of dispersal strategies including most modern categories of winged disseminules, pods, capsules, explosive dehiscence, a single arillate seed and two seed-types with dispersal hairs (one a coma). There is no direct evidence of epizoochory. In terms of mammalian frugivory the flora contains examples of all potential dietary categories. Tough and hard materials are abundant and soft material (e.g. in fleshy fruits) is common. Gut contents preserved in many birds and mammals prove that fruits and seeds played a part in vertebrate diets and borings in one seed type (Rutaspermum) indicate seed predation by weevils. No fruits or seeds show evidence of rodent gnawing. Previous quantitative studies suggesting an equable warm and humid palaeoclimate with some seasonality for Messel are supported by the newly recognised taxa. Judging from the habit of related living taxa, the vegetation appears to have been a multistratal canopy forest, including a high proportion of lianas in addition to shrubby to arborescent taxa. Herbaceous components are also present but relatively underrepresented. Among other large Eocene macrofossil floras, the Messel assemblage shows overlap with the genera known from the London Clay flora of England and the Clarno Nut Beds flora of Oregon, but relatively little similarity with floras known from eastern Asia. Compared with extant floras, the Messel flora includes a temperate component with mostly Asian endemics, and some genera that are now disjunctly distributed in the Northern Hemisphere. A large tropical-paratropical component includes genera now confined to the Old World tropics, particularly southeastern Asia and Malesia, but there are also a few exclusively Neotropical elements. Key words: angiosperms, biogeography, flora, disseminule, exceptional preservation, frugivory, lagerstätte, liana, oil shale, palaeobotany, Paleogene, Tertiary, vegetation, vertebrate diet, aril

Authors’ addresses: Prof. Dr. Margaret E. Collinson, Department of Earth Sciences, Royal Holloway University of London, Egham, Surrey, TW20 0EX, UK; Honorary Research Fellow, Department of Palaeontology, Natural History Museum London, Cromwell Road, London, SW7 5BD, UK; Corresponding Member Senckenberg Gesellschaft für Naturforschung, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; Corresponding author: Dr. Steven R. Manchester, Florida Museum of Natural History, Dickinson Hall, University of Florida, Museum Rd & Newell Dr, Gainesville FL 32611-7800, USA; Dr. habil. Volker Wilde, Sektion Paläobotanik, Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, 60325 Frankfurt am Main, Germany;

© E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), 2012, ISSN 1868–0356

Collinson, M., Manchester, S. & Wilde, V.: Fossil Fruits and Seeds of the Middle Eocene Messel biota, Germany

Introduction The oil shale of Messel near Darmstadt (Hessen, Germany) was discovered in the middle of the 19th century (Schaal & Schneider 1995). Soon after mining had started the first fossils were discovered. As a consequence of a number of papers by different specialists that were published in the 1920’s (references in Matthess 1966, Tobien 1969), Messel became a well known Lagerstätte (Tobien 1969). All of the material was collected during mining operations until scientific excavations started in the 1960’s (Kuster-Wendenburg 1969). The former oil shale mining pit of Messel is now widely known for well preserved fossils of Middle Eocene age and thus became a UNESCO World Nature Heritage Site in 1995 (Schaal 1986, 2005). Plant fossils from Messel were initially mentioned by Chelius (1886) and later monographed by Engelhardt (1922) for the first time. The posthumously published monograph deals mainly with leaves, but includes some fruits and seeds; descriptions and comparisons were based on gross morphology and venation. As a result of a revision of old leaf material, Schweitzer (in Matthess 1966: 32f) listed seven genera in six extant families of angiosperms, but the number soon increased to 24 genera in 10 families (Sturm in Tobien 1969: 173). In an extensive study including morphological and cuticular characters Sturm (1971) later focused on leaves with affinities to Lauraceae. As a consequence of increasing excavation activities by different institutions in the then abandoned and endangered pit, a great number of plant fossils were collected starting in the mid 1970’s. Stimulated by Friedemann Schaarschmidt, the new material was studied with the application of newly developed routines for preparation using ultrasonic devices under water, storage in glycerol and imaging by various methods including epifluorescence microscopy (Schaarschmidt 1982, Ackermann et al. 1992). Following an initial note on the Messel flora by Schaarschmidt (1981), pollen and spores were monographed by Thiele-Pfeiffer (1988) and leaves by Wilde (1989). Preliminary treatments of fruits and seeds were undertaken by Collinson (1982, 1986, 1988) and flowers by Schaarschmidt (1984, 1986). Goth (1990) and Lenz et al. (2007b) described different kinds of algae by thorough SEM studies. The state of the art on the flora was summarized at different stages of knowledge on the systematic composition of the plant taphocoenosis (Schaarschmidt 1988, Wilde 2004). Important information on the Messel flora is also found in a number of papers on selected taxa and organs such as leaves of Lauraceae (Sturm 1971, Kvaček 1988), seeds of Rutaceae (Müller et al. 1985, Collinson & Gregor 1988), different remains of palms (Schaarschmidt & Wilde 1986, Harley 1997), fruits of Juglandaceae (Manchester et al. 1994), leaves of Comptonia L'Hér . (Myricaceae: Wilde & F rankenhäuser

2000), wood with affinities to Buxaceae (Wilde & Süss 2001), leaves and fruits of Cedrelospermum Saporta (Ulmaceae; Wilde & Manchester 2003), leaves of Araceae (Wilde et al. 2005), fruits of Anacardium L. (Manchester et al. 2007), fruits of Cyclanthaceae (Smith et al. 2008), infructescences of Cyperaceae (Smith et al. 2009b) and leaves and fruits of Malvales (Kvaček & Wilde 2010). Additional taxa were included in a methodological paper on the application of laser scanning microscopy to flowers with in-situ pollen (Wilde & Schaarschmidt 1993), in a paper on biomarkers from a mastixioid fruit containing resin (Van Aarssen et al. 1994), and in some papers comparing different localities of similar age (Wilde 1995, Wilde & Frankenhäuser 1998, Manchester 1999) and comparing this flora with other examples of exceptional preservation (Collinson et al. 2010). Messel is the most diverse Middle Eocene fruit and seed assemblage to be documented in Europe and one of the most diverse Paleogene plant taphocoenoses worldwide. The fruit and seed flora documented here contains several new taxa, including early records of families or genera and many new morphotypes. In some cases seeds have been found in fruits and sometimes fruits are still together in infructescences. In some instances, organic attachment proves the links between different organs, e.g. leaves and fruits. Furthermore, there is the opportunity to compare the systematic diversity and composition of fruits/seeds, pollen and leaf records which have now all been studied in considerable detail (Wilde 2004). The well preserved fruit and seed taphocoenosis of Messel also provides information on dispersal biology. Gut contents may be identified and give a direct clue to the diet of the respective animals (e.g. Schaarschmidt 1992), and specific traces of herbivory may assist in systematic assignment of the host plant (Wappler et al. 2010). Messel is important for comparison with other floras of similar age elsewhere in Europe, Asia and the New World with respect to diversity, phytogeography, climate and palaeoecology; including the reconstruction of the habitat for the insects, birds and mammals. When known in sufficient detail, evidence from the Messel fruit and seed flora may also be used for calibration of modern molecular phylogenies and for modern phytogeographic studies.

Geologic setting and age The oil shale of Messel is the best known of about half a dozen isolated occurrences of Paleogene sediments from the Sprendlinger Horst (Harms 1999), the northernmost extension of the Odenwald structure flanking the northern part of the Upper Rhine Graben to the East. Except for Grube Prinz von Hessen, Paleogene sediments are confined to volcanogenic structures, and most are interpreted probably as maars formed by phreatomagmatic activity

Fig. 1: Geological setting of the Messel site near Darmstadt (State of Hessen, Germany), modified from Lenz et al. 2009.

(Felder et al. 2001). The geological context of the Messel oil shale is presented in the map and stratigraphic column of figures 1 and 2. A research drilling (core Messel 2001) in the centre of the structure at Messel penetrated the lacustrine sedimentary succession of the Messel Formation into underlying volcaniclastic deposits and, finally, vent breccias. This provided definitive proof that the Middle Eocene oil shale of Messel represents deposits of a maar lake (Schulz et al. 2002, Felder & Harms 2004) which formed soon after eruption(s) ceased. The Messel Formation was initially defined by Weber & Hofmann (1982) and then subdivided by Felder & Harms (2004) with further slight modification by Lenz et al. (2007a). Sedimentation of the Lower Messel Formation started with coarse clastic debris resulting from slope failure. With increasing stability of the slopes, individual mass-flow events may be distinguished as turbidites in a background of clay and, later, even some oil shale (Felder & Harms 2004, Lenz et al. 2007a). The Middle Messel Formation, sensu Lenz et al. (2007a), includes the typical Messel oil shale as known from strata presently exposed in the pit. It was formed under permanently meromictic conditions in the lake (Irion 1977, Goth 1990). A maximum thickness of 91.5 m of the Middle Messel Formation was preserved at the site of the research core, which is equivalent to about 640,000 years as calculated from an average sedimentation rate of 0.14 mm/yr (Lenz et al. 2011). The

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Upper Messel Formation is not known in detail since it was largely removed by mining. As seen in recent cores through a surviving area, it includes a mixed succession of organic-rich clay, silt and sand, some lignite and most probably represents the silting-up stage of the lake and/or the marginal equivalents of the Middle Messel Formation (Matthess 1966, Felder & Harms 2004). Except for microfossils such as pollen, spores, resistant remains of algae and sponge spicules, fossils have only been studied from excavations in an upper part of the Middle Messel Formation (c. 40 m according to Franzen et al. 1982) which was subject to mining and is still exposed in the present pit. During early studies the biostratigraphic age of the fossil bearing oil shale at Messel was recognised as Middle Eocene by characteristic vertebrates (Haupt 1911). This was confirmed by later studies and specified as early Middle Eocene age (Tobien 1968) or lower Geiseltalian (MP 11) in the European vertebrate chronology (Franzen 2005a, b). A lower Middle Eocene age was also obtained from palynological studies (Thiele-Pfeiffer 1988; Krutzsch 1992, SPP-Zone 14/15). The core Messel 2001 finally offered the chance for radiometric dating of the underlying volcaniclastic material (47.8 million years ago (mya); Mertz & Renne 2005). We continue to use the geochronologic term “Tertiary” as appropriate according to longstanding tradition and continuing usefulness, as defined in numerous dictionaries.

Modes of preservation The lacustrine plant taphocoenosis of the Messel oil shale is exceptional in comprising different parts and structures of plants, sometimes even in organic connection. In addition to the fruits and seeds considered here, there are leaves, pollen and spores, flowers, woody twigs and axes as well as remains of roots. Remains of angiosperms are dominant, but various conifers and pteridophytes have also been found. Algae are represented not only by resistant cysts of dinoflagellates and Zygnematales, but also by resistant sheaths and cell walls of coccal green algae like Botryococcus Kützing and Tetraedron Kützing. Remains of cell walls of Tetraedron are the major component of the organic material in the oil shale but are lost from palynological preparations due to their minute size (c. 4 µm diameter). Therefore, they may be recognised only by SEM (Goth 1990). Unlike many Eocene lacustrine deposits in which the fruits are preserved as impressions, e.g. Green River, McAbee, Republic, in western North America, most of the plant fossils in the oil shale at Messel are preserved as remnants of the original organic material in various stages of compression and degradation. Although compression of the fossils in the sediment has resulted in distortion, i.e.


Fig. 2: Stratigraphy of the Messel Formation at Messel (modified from Lenz et al. 2011); a: Lithologic succession of research core Messel 2001; b: Stratigraphic position of the Middle Messel Formation slightly modified from Mertz & Renne (2005).

flattening perpendicular to the pressure, the tissues and anatomy are often well-preserved, including some more labile tissues as well as the sclerified parts. The degree of compression mainly depends on the rigidity of the original tissues. Three-dimensional preservation of cellular structures is sometimes seen in sclerenchymatous tissues, especially in fruits and seeds. Woody tissues may be found well preserved and in different stages of compression and homogenization, even within the same specimen (Wilde & Süss 2001). Most of the leaf material is compressed, sometimes even degraded except for the network of the venation; three dimensional preservation of leaf parenchyma is a rare exception. Occasionally, individual leaves and a few fruits [e.g. one Cyclanthus Poit. fruit, Smith et al. (2008)] are preserved as, or covered by, a powdery film of pyrite. Preservation of cuticles is fairly frequent, although its quality is different from taxon to taxon (Wilde 1989). Due to a low maturation of the organic material (mean vitrinite reflectance 0.26 %; Rullkötter et al. 1988), the

oil shale at Messel is suitable for biomarker studies (e.g. Mattern et al. 1970, Habermehl & Hundrieser 1983a, b, Goth et al. 1988, Robinson et al. 1989, Van Aarssen et al. 1994, Adam et al. 2006).

Material and methods Specimens are recovered during the same excavations from which animal fossils are obtained. Large slabs of the oil shale are removed from the quarry face. They are split in the field by knives and the fresh surfaces are immediately checked for fossils by eye, if necessary assisted by hand lens observation. To prevent fracturing, that occurs on drying due to differential contraction of the shale and organic material, specimens are immersed in water immediately after collection. Generally the specimens are retained on the original shale slabs and these are carefully trimmed around the fossils if necessary.

For further treatment plant fossils are transported to the Senckenberg laboratories in Frankfurt. They are prepared, under a stereomicroscope while immersed in water, with a handheld dental sonication tool (DentsplyCavitron Model 700-II B) to remove adhering shale from the exposed surfaces of plant material including leaves, flowers, fruits and seeds (Schaarschmidt 1982). For delicate specimens fine needles are used for carefully scratching away the sedimentary cover. In some cases, the seeds or fruits were physically dislodged from the sediment to reveal all external surfaces. Specimens are stored in glycerol to prevent desiccation with the addition of thymol as a fungicide (Schaarschmidt 1982). A few specimens from old collections are stored dry. Light macrophotography was performed using digital cameras (Nikon Coolpix 995, Nikon D-300; Canon Powershot A640), mounted on a copy stand with a pair of strong incandescent side lights adjusted for oblique lighting to highlight surface details. Light microphotography was done with the Nikon camera mounted on a Zeiss M400, or, in some instances, with the epifluorescence apparatus described by Schaarschmidt (1982). For photography, specimens were usually removed temporarily from the glycerol in which they are stored, and placed in a tray of water. In other cases, superior results were obtained by allowing the specimen surfaces to begin drying, and photographing specimens without liquid. For electron microscopy, specimens were cleaned of adherent siliciclastics with a brief treatment in HF, neutralized, and dried naturally in air. For SEM they were then mounted on stubs using glue (Bostik; Bostik Ltd, Leicester, UK) on a cover glass or self-adhesive pads. Some specimens were coated with gold in a Polaron sputter coater and examined using a Cambridge S600 and a Philips 501B SEM (both now decommissioned; formerly at the Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, and King’s College, London, respectively). Other specimens were mounted using glue (Bostik) on a coverglass, sputter coated with gold in a Polaron (Quorum Technologies Ltd., Ringmer, UK) E1500 sputter coater, and examined in either an Hitachi (Hitachi High Technologies Ltd., Tokyo, Japan) S3500N SEM or a FEI (FEI Company, Hillsboro, Oregon, USA) Quanta 200F field emission ESEM. For TEM, specimens were embedded in Spurr resin (Agar Scientific Ltd., Stanstead, UK). Sections 60 nm thick were cut with a Diatome (Diamond AG, Biel, Switzerland) diamond knife using a Reichert Ultracut E ultramicrotome (Leica Microsystems, GmbH, Wetzlar, Germany) and examined unstained using a Hitachi H-7600 TEM (Hitachi High Technologies Corporation, Tokyo, Japan). Epifluorescence light microscopy (LM) imaging was initially promoted by Friedrich & Schaarschmidt (1977) for plant fossils and later especially adapted for material from Messel by Friedemann Schaarschmidt (Schaarschmidt 1982). This technique takes advantage of pollen and cuticle autofluorescence to enable imaging


of pollen and cuticles in situ without chemical preparation, thus avoiding any damage to the fossils. The selfmade macrofluorescence system is now dismantled since commercial systems have been developed. The light source was a mercury high pressure lamp (HB0 50) with a Schottglas BG 3 exciter filter and a Kodak Wratten number 12 barrier filter. Colour images were captured on Ektachrome or Fuji 400ASA film and black and white images on Kodak Tri-X Pan 400 ASA film exposed at 800 ASA and developed in Promicrol. Microfluorescence observation and imaging was undertaken using the same films and the same light source on a Leitz Dialux microscope with a 440 nm exciter filter, a 460 nm beam splitter and a 475 nm barrier filter. Other microfluorescence observation and imaging (as used in Smith et al. 2009a, b) was undertaken on a Zeiss (Carl Zeiss, Oberkochen, Germany) standard laboratory microscope with a 436 nm exciter filter, a 460 nm beam splitter and a 470 nm barrier filter (blue-violet filter set) using a mercury high pressure lamp (HB0 50) light source in a Zeiss epifluorescence condenser IV FI. Color images were captured on Kodak (Eastman Kodak Co., Rochester, New York, USA) Ektachrome 400ASA slide film. Attempts to embed specimens in carbowax (MEC pers. obs.) or standard resins (Smith et al. 2009b) for sectioning were not successful as the embedding media did not penetrate through the specimens. With the advent of new technology, a few specimens have been studied recently by Synchrotron Radiation X-Ray Tomographic Microscopy (SRXTM) and one or two by Computed Tomography (microCT). Some have been found to reveal excellent details of internal morphology and anatomy by SRXTM (Smith et al. 2009b; and some illustrations herein) and we envisage a future long term project applying these methodologies to Messel fruits and seeds. SRXTM imaging was performed at the TOMCAT beamline (Stampanoni et al. 2006), Swiss Light Source, Paul Scherrer Institute, Villigen, Switzerland to reveal details of internal structure. Oil shale blocks with specimens were trimmed to a suitable size, rinsed and briefly soaked in water to remove glycerol. Specimens were imaged with the small blocks of oil shale either (i) immobilized at the base (and sometimes also at the sides and top) with foam (plastzoteTM) in a plastic tube or (ii) held in a metal clamp. Specimens released from oil shale were placed in a “cocoon” of plastzote. Data were acquired using the 1.25, 2 × or 4 × objectives on an X-ray microscope, and an exposure time 180–700 ms at 19.8 keV for Messel specimens still in oil shale and 10 keV for specimens clear of oil shale. A total of 1501–2048 projections were acquired over 180°. Projection data were processed and corrected sinograms were then used for reconstruction. MicroCT imaging was undertaken at the Natural History Museum, London, UK on a Metris X-Tek HMX-ST 225 scanner using a tungsten reflectance target, a voltage of either 105 or 180 kV, a current of either 71 or 190 mA and an exposure time of 250 ms. Due to the longer scan


time (c. 1 hour compared to c. 10 mins) specimens were wrapped in cling film to reduce risk of damage by drying. Oil shale blocks were inserted into small blocks of Oasis TM (foam used in flower arranging) to hold and support them. Reconstructed images from SRXTM and microCT were processed using Avizo 5.1 (Mercury Computer Systems, France). Images were captured in Avizo itself and then processed in Adobe Photoshop CS for Mac OS X. For measurements accompanying the descriptions of species, we have stated the number of specimens studied when the number is small (3 or fewer) and might be unreliable as to the typical variability. Observations are by standard light microscopy, unless otherwise indicated in instances where scanning electron microscopy (SEM), epifluorescence, SRXTM or microCT were used. Scales in LM photographs have smallest units in mm (each mm being measured from mid-point to mid-point of the black calibration lines). We studied plant fossils from Messel mainly stored at the Senckenberg Forschungsinstitut and Naturmuseum in Frankfurt am Main (specimens numbers with prefix SM.B Me), but also used the less extensive material of the Hessisches Landesmuseum Darmstadt (prefix HLMD-Me-). Comparative work on extant fruits and seeds was undertaken at the herbaria of Harvard University, Missouri Botanical Garden Herbarium, the fruit and seed collection of the Humboldt Museum Berlin; and the US National Fruit and Seed Herbarium, Beltsville, MD. Soft tissues were removed after hydration with boiling water to expose the endocarp or seed. Comparisons with fossils were made at the Natural History Museum, London (BMNH; now NHMUK), the Palaeobotanical Herbarium, Institute of Botany, Chinese Academy of Sciences, Beijing (CMPH), the Field Museum, Chicago (FMNH), the Geological Institute, Russian Academy of Sciences, Moscow (GINRAS), the Palaeobotanical Museum of the W. Szafer Institute of Botany, Polish Academy of Sciences, Krakow, Poland (KRAM-P), the Florida Museum of Natural History, University of Florida, Gainesville, FL(UF), the United States National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM), the Burke Museum of Natural History and Culture, University of Washington, Seattle (UWBM), the Denver Museum of Nature and Science (DMNH), the Naturhistorisches Museum Mainz/Landessammlungen für Naturkunde RheinlandPfalz, and the Humboldt Museum, Berlin (MfN). With a goal of completing this survey in a timely fashion and keeping in mind that the collections at Senckenberg Forschungsinstitut und Naturmuseum are extensive and new specimens are being added all the time, we have not been able to deal with every specimen of fruit or seed collected from Messel. To the best of our ability we have documented elements with distinctive morphologies added to the collections before the end of 2010. This treatment may have overlooked some additional taxa whose visible external morphology is not distinctive. Xray technologies may prove useful for the study of these in the future. Very small seeds of a size class less than two millimeters have likely been overlooked because the

techniques employed in the field were directed mainly towards recovery of megafossil remains. This monograph deals only with fruits and seeds from the Messel oil shale. In a few cases (e.g. Toricellia DC. and Rutaceae) we refer to specimens recovered by sieving from the clastic sequence at site SMF 7. These specimens were studied by Collinson (1988) and Collinson & Gregor (1988); details of the material and site can be found in those papers.

Floristic composition In this compilation, we recognise 156 morphologically distinct species or morphotypes. Ninety-one of these species representing 75 genera are assignable to modern families, but an additional 65 species remain uncertain as to their position relative to extant families. The latter, placed in the fossil genera Saportaspermum Meyer & Manchester, Spirellea Knobloch & Mai, and Carpolithus L., probably include extinct genera as well as those whose identification requires still more detailed comparative work. The fruit and seed assemblage is dominated by angiosperms, although the conifer Doliostrobus Marion is also present. Table 1 (see page 86) presents the list of taxonomically determined fruits and seeds organized in the phylogenetic sequence of APG III (Angiosperm Phylogeny Group 2009), although with some modification because we accept the cornalean families Mastixiaceae and Nyssaceae as distinct from Cornaceae. The most diverse families are Menispermaceae (17 morphotypes), Icacinaceae (11 morphotypes), Vitaceae (7 morphotypes), and Mastixiaceae (5 morphotypes), followed by Anacardiaceae (4 morphotypes), Rutaceae (5 morphotypes) and Juglandaceae (3 morphotypes). Collinson (1988) documented the relative abundance of fruit and seed taxa (n = 3640) at SFN in April 1987 when, according to Schaarschmidt (verbal communication 1986), most specimens found in the field were collected and prepared. The most abundant taxa were Friedemannia gen. n. (41 %), Euphorbiotheca Reid & Chandler (5.5 %), Volkeria Smith, Collinson, Simpson, Rudall, Marone & Stampanoni (3.5 %) and Eomastixia Chandler (3.2 %) (treated as Palmae, Euphorbiaceae, Cyperaceae, and large ovoid fruit with resin, respectively, in Collinson 1988). In the same study, members of the Vitaceae, Icacinaceae, Rutaceae, Hamamelidaceae, Carpolithus sp. 42 (listed as “ovoid shiny drupe”) Carpolithus sp. 57 (seed from the Masillamys gut) had percentage occurrences of 2 to 3.2 %. However, we do not have formal abundance records of taxa in the field and the relative abundances of taxa in the current museum collections do not reflect field abundance because subsequently priority was placed on retention of rare and unique specimens in addition to adequate representation

of the abundant taxa. Nevertheless, it is still clear from field experience (VW) that the by far most common taxa include Friedemannia and Euphorbiotheca. Only 60 % (93/156) of the Messel fruit and seed taxa are placed with confidence in modern families. This reflects our philosophy that confident identification requires recognition of multiple diagnostic characters – a combination of characters confined to a particular extant genus or family. We tried to err on the side of caution – placing taxa in incertae sedis (e.g. Carpolithus spp.), rather than in similar modern taxa when evidence was not considered overwhelming. In the discussion of unplaced taxa, we provide comments that might be helpful in the continuing search for systematic placement of these fossils, but we caution against the use of this information for calibrating molecular phylogeny or for climatic interferences, unless more detailed investigations are carried out to confirm or dismiss these suggestions. Among the species with secure familial position, some are clearly belonging to extinct genera, falling outside the morphological range of the extant genera, e.g. Cedrelospermum, Cruciptera Manchester, Eomastixia, Friedemannia, Hooleya Reid & Chandler, Karinschmidtia gen. n., Martinmuellera gen. n., Palaeophytocrene Reid & Chandler, Volkeria. Others fit modern genera, like Ailanthus Desf., Alangium Lam., Anacardium L., Cyclanthus, Lannea A. Rich. in Guillem., Toricellia, Meliosma Blume, and Vitis L. Still others are placed in fossil morphogenera that accommodate morphology found in more than one modern genus, as in the case of Euphorbiotheca, Laurocarpum Reid & Chandler, Mimosites Bowerbank, Leguminocarpon Goeppert, Myristicacarpum Gregor, and Rutaspermum Chandler. Genera that are newly named in this monograph include Friedemannia, Karinschmidtia, Martinmuellera and Camelliacarpoidea. However, it is likely that most of the 62 morphotypes attributed to Carpolithus are distinct enough to be placed in separate genera. Some taxa that might have been expected are apparently missing from this assemblage, including families like Platanaceae, Fagaceae, and Betulaceae, but they are known from leaves and/or pollen. For further discussion see next section.


and/or seeds (table 2. see page 88). Previous leaf records of Engelhardt (1922) of Magnoliaceae, Rhamnaceae and Simaroubaceae are purely based on macromorphological characters and therefore not sufficiently reliable, and the type of Catalpa spiegelii Engelhardt 1922 (putative Bignoniaceae) has recently been transferred to the Malvaceae (Kvaček & Wilde 2010). Four families are known from both leaves and pollen (Thiele-Pfeiffer 1988, Wilde 1989), but are absent in the confirmed fruit and seed record: Araliaceae, Ericaceae, Myricaceae and Symplocaceae. There is some potential for Ericaceae fruits to be present in Carpolithus species and the Messel collections are probably biased against small seeds such as those typical of some Ericaceae. However, the characteristic and well known fruits of the Symplocaceae have definitely not yet been identified in spite of pollen and a putative leaf record. Betulaceae are present in the pollen record (Thiele-Pfeiffer 1988) and most probably also represented among unpublished leaf records (VW, personal observation) but are not present amongst the fruits and seeds. Pinaceae and Fagaceae are well represented in the pollen record, but have not yet been identified among more than 25,000 plant macrofossils in the collections. This has been interpreted in terms of the probability that they grew some distance from the lake, outside of the immediate catchment area (e.g. Wilde 1989, 2004). Only a single moderately preserved leaf may be referred to Platanaceae (Wilde 1989) which are not known from pollen or fruits/seeds. Platanaceae are therefore expected to have been extremely rare around the lake at Messel. If the macrofossil record (leaves, fruits, seeds, flowers and wood) is pooled together with the palynorecord at the present state of our knowledge, there is a cumulative diversity of 75 families (11 of them monocotyledons) and 203 “species” (16 of them monocotyledons) of angiosperms which have been described or mentioned from the oil shale of Messel in more modern studies (Sturm in Tobien 1969; Sturm 1978; Collinson 1982, 1986, 1988; Thiele-Pfeiffer 1988; Wilde 1989, 2004; Kvaček & Wilde 2010, this paper). Since many morphotaxa of leaves, fruits, seeds, flowers and pollen are known (not yet identified to modern families) and some remain to be described, the true diversity of angiosperms must have been considerably higher.

Comparison with diversity known from leaves and pollen In the present monograph, 34 families of angiosperms are identified from Messel with 140 genera (counting 9 Carpolithus representatives as separate genera). Three families and three genera represent monocotyledons. Thirteen of the dicotyledonous families (Alangiaceae, Altingiaceae, Bignoniaceae, Burseraceae, Elaeocarpaceae, Euphorbiaceae, Magnoliaceae, Myristicaceae, Rhamnaceae, Sabiaceae, Simaroubaceae, Tapisciaceae, Toricelliaceae) and the mococotyledonous Cyclanthaceae are only known from fruits

Fruit and seed biology, dispersal and animal diets

Soft to leathery fleshy fruit tissues A large number of Messel taxa have the entire fruit wall (pericarp) organically preserved (table 3, see page 90).


This not only provides conclusive proof of the nature of fruit tissues but also helps to evaluate inferences made from nearest living relatives for less exceptional organic preservation states (in other fossil floras) that may only have endocarps or seeds preserved. Some shrinkage may have occurred during the fossilisation process and most specimens are strongly compressed which makes it inappropriate to estimate propagule volumes. However, measurements of fruit wall layers are likely to be reasonably close to (or possibly slightly greater than) their original thickness as they are mostly measured at the sides of specimens and there are no obvious large compression rims. Furthermore there is minimal evidence of fruits splitting or bursting during compaction. A number of fruits clearly possessed soft or fleshy tissues proven by (i) preservation of epicarp with space between that and endocarp, (ii) the preservation of parenchymatous tissue, (iii) crumpling or wrinkling of inner fruit tissues or epicarp cuticles. Examples are found in the Vitaceae, Menispermaceae, Lauraceae, and several Carpolithus species (table 3). In many cases softer mesocarp tissues surrounding endocarps rarely exceed 2 mm in thickness and so would provide only a small amount of more digestible tissues for animal consumption. Exceptions include C. sp. 18 a large fleshy fruit up to 6 cm in diameter; other larger fruits with soft tissues e.g. C. sp. 15, C. sp. 19, C. sp. 46; several berries including C. sp. 6, C. sp. 30 and C. sp. 53 the latter of which we infer possessed extremely soft fleshy tissues, the softest in the flora (table 3). In addition there is the presence of a distinctive component such as the hypocarp in Anacardium (Manchester et al. 2007). Whereas modern Anacardium are often dispersed by bats, none of the Messel bats were frugivorous based on their gut contents (Storch 2001, Storch et al. 2002) and Eocene bats in general are considered to be insectivorous (Simmons et al. 2008). Therefore other frugivores presumably fed upon and possibly dispersed Messel Anacardium (Manchester et al. 2007) and other Messel fleshy fruits.

Vertebrate gut contents containing fruits and seeds Vitaceae seeds are present in the gut content of the Perissodactyl Eurohippus parvulus (Laurillard) (formerly Propalaeotherium parvulum Laurillard, revised by Franzen 2006). These were illustrated by Koenigswald & Schaarschmidt (1983). However, seeds of Vitaceae are present in only one of the 44 individuals examined by Franzen (in Storch et al. 2005) and most gut contents are leaf fragments (Franzen 1992, Franzen in Storch et al. 2005, Franzen 2007a, Wilde & Hellmund 2010). Wilde & Hellmund (2010) note, based on Franzen (2007b: pl. 26), that a single Propalaeotherium hassiacum Haupt specimen also contains Vitaceae seeds (n = 15, Franzen in Storch et al. 2005). As these seeds are intact in the gut (Koenigswald & Schaarschmidt 1983: fig 5) the perissodactyl would have acted as a dispersal agent for the seeds (endozoo

chory). These hoofed, ground-dwelling mammals could not have sourced these fruits high in the canopy but likely obtained them as fallen fruits on the ground. Based on the illustrations in Koenigswald & Schaarschmidt (1983) the vitaceous seeds are small (c. 4–6 mm long and c. 3 mm wide) with a centrally positioned chalazal scar, from which a groove extends to seed apex and base, and a more or less smooth dorsal seed surface with faint radial striations. The ventral grooves are long but their angle of divergence is obscured by the angle of compaction. These seeds are closest to those identified as Parthenocissus britannica (Heer) Chandler in the present work. Seeds of Carpolithus sp. 57 are present in the gut content of one specimen of the rodent Masillamys Tobien (MEC and G. Storch unpublished data, Collinson 1988). The gut of this specimen is completely filled with seed fragments which, when examined by SEM (Collinson 1988), have identical anatomy to C. sp. 57 (herein pl. 73). This anatomy is diagnostic to this taxon at Messel and the seeds were small and brittle hence easily crushed. The very shiny surface may indicate their former presence in a soft fruit, or possibly former presence of an aril, but all the oil shale specimens are isolated seeds. Unfortunately, the identification of this seed type has so far eluded our efforts (unpublished work by S. Y. Smith and MEC) but it is one of the most abundant small-seeded taxa in the collections. According to Franzen (in Storch et al. 2005) Masillamys were small rodents that led a simple terrestrial life on the forest floor lacking any adaptations for life in the trees or for digging. The fruit containing these seeds may therefore have been fallen fruit or may have been borne by low-growing plants. As the seeds are all fragmentary in the gut at least this individual of Masillamys acted as a seed predator, not a seed disperser. Mammals and birds may have been involved in the dispersal of the arillate seed (see section on insect dispersal). Morlo et al. (2004) provide a listing of Messel vertebrates with 45 mammal species and 43 bird species from which references to detailed taxonomic treatments can be obtained. Fruits and seeds in gut contents, other than those discussed above, are reported for a number of Messel mammals and birds but none of the fruits and seeds have been identified, either to a plant group or to a taxon present in the oil shale. Other mammals with fruits and seeds in the gut include the artiodactyls Messelobunodon Franzen, Eurodexis Erfurt & Sudre and Aumelasia Sudre, the paroxyclaenid Kopidodon Weitzel, the pantolestid Buxolestes Jaeger, the rodent Eogliravus Hartenberger (Franzen & Richter 1992, Koenigswald & Storch 1992, Koenigswald et al. 1992, Franzen in Storch et al. 2005); and the primate Darwinius Franzen et al. (Franzen & Wilde 2003, Franzen et al. 2009). Birds with fruits and seeds in their gut contents include the gruiform Messelornis Hesse (Morlo 2004), a piciform bird (Peters 1992: fig. 217), two coraciiforms (rollers) Primobucco Brodkorb (two specimens with accumulations of large >1cm seeds) and Eocoracias


Mayr & Mourer-Chauviré (one seed in one specimen) (Mayr et al. 2004), two species of Primozygodactylus Mayr (Zygodactylidae), small arboreal birds, (Mayr & Zelenkov 2009) and the colliiform stem group sandcoleid Eoglaucidium Fischer (Mayr 2009). In this last case the single large seed c. 1.3 cm long was suggested to be “probably from a species of the Annonaceae (custard apples)” (Mayr 2009: fig. 16.2, tentatively determined by VW). We have not proven any Annonaceae from Messel and so it would be important to re-examine this specimen to see if the seed determination can be confirmed. The characteristic ruminate endosperm of Annonaceae is not evident on the illustration (Mayr 2009: fig. 16.2) but it is possible that this pattern may not always be expressed on the outside of the seed coat. As discussed by Mayr (2005) some Paleogene birds show different dietary and/or locomotor adaptations to the living members of their clades. Therefore, the Messel gut content records of fruits and seeds are important evidence of ancient diet. Equally, amongst the mammals, it was unexpected to find fruits in the gut content of the piscivore Buxolestes Jaeger (Koenigswald et al. 1992). Hopefully the production of this monograph will be a first step to identifying more of these gut content seeds, although curatorial restrictions still remain in that destructive analysis (e.g. preparation of fragments for SEM) may not be allowed for rare, skeletal specimens and CT analysis is unlikely to provide the required resolution of fruit wall and seed coat anatomy.

Insect dispersal and feeding Borings in numerous Rutaspermum chandlerae Collinson & Gregor seeds from site SMF 7 cannot be distinguished from those made in grain by modern weevils (Collinson & Gregor 1988; Collinson 1990, 1999a, b). Similar borings occur in the same seed type elsewhere in Europe and in North America at other middle Eocene sites but not in older strata (Collinson 1999a, b; Hooker & Collinson 2001). Weevils occur in the Messel oil shale (Lutz 1992; Wedmann 2005a, b), and represent a high percentage of the beetle fauna, but no detailed study has yet been published. The Rutaceae are represented by only a small number of specimens in the oil shale (see systematic part), nevertheless one specimen of Rutaspermum messelense Collinson & Gregor has a boring like those in R. chandlerae. There is a single arillate seed in the Messel fruit and seed flora, Carpolithus sp. 58 (pl. 78). Presence of an elaiosome (a soft lipid- or oil-rich seed appendage, including some arils) is the signal adaptation for ant dispersal (Lengyel et al. 2010). Ants are present in the Messel insect fauna (Lutz 1992; Wedmann 2005a, b) but we are not aware of any studies on their potential roles as seed dispersers. According to the extensive survey by Lengyel et al. (2010) at least 77 families and 334 genera of flowering plants (both monocots and dicots) include species with ant

dispersal (myrmecochory). Although it is clear that the aril on C. sp. 58 was very compact (not loose hairs) we do not know if it was lipid-rich and we do not know the original colour, both of which are important for understanding likely potential dispersal agents for arillate seeds. Bach et al. (2007) noted that birds were important dispersers of lipid-rich arillate seeds whilst Garcia-Robledo & Kuprewicz (2009) and Christianini & Oliveira (2010) both report complex interactions involving mammals, birds and ants in dispersing lipid-rich arillate seeds (including those with red or orange colours often thought to imply bird dispersal). Therefore, although it seems likely that ants were involved at some point in dispersing this seed type (as primary or secondary seed dispersers or as seed rescuers), we are unable to confirm the dispersal biology. To our knowledge this seed may be the earliest in the fossil record to prove the presence of an aril through exceptional organic preservation. Interactions between the fruits and seeds and insects at Messel are very limited, especially compared to those with vertebrates. Although there may well be other rare specimens that show evidence of insect interaction present in the fruit and seed collections these are the only two noteworthy examples that we have observed.

Dry fruits and seeds: dehiscence mechanisms and vertebrate diet Many Messel taxa produced dry fruits including capsules and valvate fruits and pods (table 3). One of the pods shows evidence of dehiscence and it is reasonable to infer this for others based on living relatives. Capsules and valvate fruits show clear evidence of splitting to release dry seeds. The Messel flora includes some examples of possible explosive dehiscence such as Euphorbiotheca with variously twisted endocarps (pls 10, 11) sometimes partly to completely detached from the surrounding mesocarp. Torpedo shaped dry seeds with prominent hilar scars (pl. 14f–t), and fruits of Hamamelidoideae (pls 13, 14) appear to show the syndrome of explosive seed dispersal known from the extant genera. Many taxa at Messel produced dry seeds or endocarps (table 3) a suitable resource for seed feeders (including mammals or birds). Many of these taxa have thick, hard and tough seed coats and other taxa have thick, hard, tough endocarp walls hence their food resources would be suitable for mammals able to gnaw to gain access. To our knowledge the earliest example of rodent gnawing is that documented by Collinson & Hooker (2000) from the late Eocene and we have not seen any Messel fruits or seeds showing evidence of mammalian gnawing.

Dry fruits and seeds as a vertebrate food resource: fibrous, tough, woody and resinous tissues The Messel flora contains one or two large seeds (e.g. Sapotispermum and C. sp. 18) but most seeds are smaller,


e.g. 4–8 mm in maximum dimension. The food resources within each seed would therefore have been relatively small (by comparison with larger modern nuts such as Corylus L. or Juglans L. for example) so larger numbers would need to be consumed for an equivalent volume of internal tissues. In the absence of evidence of gnawing (see above) the hard and tough seeds and endocarps, which formed parts of animal diets, were presumably chewed (i.e. processed in the mouth) by mammals to access their internal food reserve. Mesowear and microwear are important features that are produced by chewing and can be observed on fossil teeth as indicators of diet in Paleogene mammals (Joomun et al. 2008, 2010). Characteristics of food (including fruits and seeds) that can influence mesowear and microwear include (i) abrasive material (such as phytoliths); (ii) soft material which yields readily to pressure (such as fleshy soft fruit tissues); (iii) tough material (toughness relating to resistance to crack growth) which is difficult to break down (such as woody sclerotic tissue of seed coats and endocarps); (iv) hard material which is resistant to deformation under indentation (such as seed coats and endocarp walls) and (v) fibrous material which has a structure composed of threads (summarised from Joomun et al. 2010). Soft material has been discussed above. Phytoliths are unlikely to be preserved at Messel as the oil shale is decalcified and even siliceous diatoms are only represented by moulds. A few seeds show spaces in outer layers of seed coats that might originally have contained crystals (e.g. Carpolithus sp. 14, pl. 52e, f; C. sp. 34, pl. 60h–l; C. sp. 57, pl. 73o). These possible crystal cells are best observed by SEM of fractured seed coats, and not all taxa have yet been examined in this way. Nevertheless, the evidence suggests that such crystal cells are rare in the Messel fruits and seeds. Several Messel fruits are fibrous (i.e. have elongate fibres visible within their tissues (pl. 2g, pl. 5a–h, pl. 28a, k–m, pl. 48a, b). These include the palm Friedemannia, some of the mastixioids, Lannea of the Anacardiaceae and C. sp. 3 (table 3). Others have closely packed elongate hair-like tissues (e.g. Sloanea L. and C. sp. 38) which may be similar to fibres in dietary terms. There are very few large woody fruits (e.g. C. sp. 54, which is the largest, and C. sp. 37). Some fruits contain resin (e.g. some mastixioids, (pls 28, 29) and C. sp. 38 (pl. 63) and one (C. sp. 37) contains preserved viscous cream-coloured material (pl. 62f, h), possibly gum-like fluid endosperm. Dry seeds include those with very brittle seed coats (that shatter rather than bend during fossilisation) such as C. sp. 18 and C. sp. 57. All dry seeds and all endocarps (table 2) have strengthened or thickened sclerotic tissue layers (see systematic part for specific details of anatomy). At least 50 % of the flora has tough or hard layers that represent a substantial proportion of the fruit or seed (relative to overall size). Therefore, in terms of mammalian frugivorous diet, the Messel fruit and seed flora contains examples of all potential dietary categories. Many fruits and seeds contain substantial thicknesses of 10

tough and hard materials. Soft material is common with significant amounts proven in a number of taxa. Fibrous material is relatively rare and abrasive material is rare.

Endosperm food reserves Two fruits from Messel have preserved ruminate endosperm, namely Myristicacarpum, (Myristicaceae) and Carpolithus sp. 24. The preservation provides proof of the presence of a well-developed endosperm food reserve but both are very rare at Messel. Annonaceae seeds are another example with ruminate endosperm which are very well represented in Paleogene floras (Collinson & van Bergen 2004) but seeds of this family have not been verified at Messel. Other examples of well-developed endosperm reserves can be inferred in seeds of Vitaceae as this is proven in permineralisations in the London Clay and Clarno Nut Beds floras (e.g. Manchester 1994). C. sp. 37 is unique in containing preserved soft viscous cream-coloured material (pl. 62f, h), possibly gum-like fluid endosperm or an endosperm decomposition product. This too indicates a copious food reserve.

Epizoochorous dispersal None of the fruits and seeds from Messel possesses hooks or barbs. A single species, Carpolithus sp. 22, (pl. 56a–k) possesses fairly long spines but their morphology is more suggestive of a function for protection (or perhaps anchorage after dispersal) than for attachment to animals. C. sp. 15 (pl. 53a–d) possesses short blunt spine-like tubercles but again these are not suited for epizoochory.

Wind dispersal There are a wide variety (but comparatively small numbers) of winged fruits in the Messel flora. These include helicopters, fin-wings, trilobed wings, bilateral wings, single-sided wings, an encircling membranous wing and a stiff circular wing (table 3). These fruits belong to the Juglandaceae, Ulmaceae, Ailanthus and Carpolithus species 20, 39 and 55. In addition there are two types of winged seeds (Saportaspermum and the very small Carpolithus sp. 21), and one fruit containing winged seeds (C. sp. 5) (table 3). This variety is consistent with that known in other Paleogene floras which encompasses the vast majority of modern categories (Collinson & van Bergen 2004, Manchester & O’Leary 2010, Wilde & Frankenhäuser 2010) which can exploit a wide variety of trajectories, attitudes and flight paths during dispersal (Burrows 1986). Carpolithus sp. 62 (pl. 76a–d) is a single specimen of a new winged fruit morphology that has not to our knowledge previously been recorded in the Paleogene. It has two long, vertically oriented wings aris-


ing from near the fruit base. There is some similarity in wing number, length and position to fruits of some modern Dipterocarpaceae [which are the most striking group of modern plants with sepaline fruit wings (Ridley 1930)] and also Gyrocarpus Jacq. of the Hernandiaceae (Ridley 1930). The wings on these fruits have only a small role in increasing dispersal distance by wind in forests but may also act like sails for water, even moving fruits upstream against currents during strong winds (Ridley 1930). However, the crumpling and folding of the wings on the Messel fruits indicates that they may have been flexible, not rigid, so these analogies may not be appropriate. One seed type, Cypselites Heer (Apocynaceae), possesses a terminal tuft or plume of hairs (coma) (pl. 4). One very small seed (seed body 1.3 mm) (Carpolithus sp. 26) has spreading fine filamentous hairs (pl. 56q–s). C. sp. 26 is not known to us from other Paleogene floras. In contrast, the coma-bearing seed Cypselites is well known and widespread in the Paleogene occurring in many floras (reviewed in Collinson & van Bergen 2004). The coma probably functioned to aid seed orientation on landing. Plumed seeds are rare in Paleogene floras in general (Collinson & van Bergen 2004). The fact that only two types have been found at Messel, where hair preservation would be favoured by exceptional preservation conditions, suggests that this rarity is a genuine reflection of the Paleogene evolutionary grade of plumed propagules. There are very few small dry seeds at Messel (e.g. those within Cyclanthus lakensis (Chandler) Smith, Collinson & Rudall or the small fruits of Volkeria (Smith et al. 2008, 2009b). Dust seeds are known in other Paleogene floras, although there are rather few examples (Collinson & van Bergen 2004). Dust seeds in the Eocene London Clay and Clarno Nut Beds floras include seeds in fruits of Lythraceae, Hydrangeaceae and Flacourtiaceae (Manchester 1994). In addition Rhododendron L. seeds are known from the Paleocene/Eocene transitional strata in the UK (Collinson & Cleal 2001c) and the late Eocene of California (Wang & Tiffney 2001). None of these are recorded at Messel. However, it is likely that isolated dispersed small seeds (e.g. below 2 mm in size) are biased against in the Messel flora (due to collecting strategy of examining bedding surfaces for macrofossils). In addition, small seeds may be present within some fruits that show no external diagnostic characters and have not yet been examined in detail.

Other dispersal mechanisms A single awned fruit is present in the flora Carpolithus sp. 2. The fruit is similar to those referred to the genus Clematis L. in the latest Eocene Insect Limestone flora of England (reviewed in Collinson et al. 2010) but differs in detail. The affinity of either the Messel Oil Shale or Insect Limestone specimens to the genus Clematis needs to be reassessed including a comparative survey of mod-

ern awned fruits. Awns may function to reduce speed of descent, alter trajectory or aid orientation on landing. Cyclanthus Poit. ex A. Rich. (Smith et al. 2008) has fruiting cycles (pl. 8a, b) and is extremely unusual in that these cycles slide down and fall off the mature fruiting axis, hence the hole at their centre (pl. 8a). Seeds are then released. For further discussion see Smith et al. (2008).

Dormancy and germination Collinson & van Bergen (2004) reviewed direct evidence in the Paleogene fossil record for endogenous (broadly equal to embryo controlled) and exogenous (seed coat imposed or enhanced) dormancy (sensu Baskin & Baskin 1998). In contrast to evidence reviewed for Paleogene floras in Collinson & van Bergen (2004) we have not identified any embryo or seedling preservation at Messel. Therefore we are not able to add to that previous review regarding Paleogene endogenous dormancy. Hard seed coats or endocarp walls impermeable to water and specialist opening mechanisms are associated with physical exogenous dormancy (Baskin & Baskin 1998). In Paleogene floras these include Anacardiaceae, Mastixiaceae and Nyssaceae fruits and nymphaealean seeds (Collinson & van Bergen 2004) along with Toricelliaceae, all of which occur at Messel (table 3). Many taxa in the Messel flora (see anatomical details in systematic section) have thick and tough seed coats or endocarp walls that may have contributed to physical dormancy.

Taphonomic Considerations Due to its phreatomagmatic origin, the Middle Eocene maar lake at Messel and its immediate surroundings most probably represented a more or less closed hydrological system. As a consequence, most of the macroscopic plant material should have been transported by runoff, minor tributaries or airborne transport into the lake from its limited catchment area (Wilde 1989, Ferguson 1993). In addition, input of fruits and seeds from feces (endozoochory) of birds and mammals must be regarded as possible (Ferguson 1993, Collinson et al. 2010, herein section on dispersal). Unequivocal adaptations for epizoochory have not been observed (Collinson et al. 2010, herein). Accidental transport by vertebrates is also possible. Airborne transport by wind or insect vectors may have contributed significantly to the pollen record in the oil shale (Ferguson 1993). However, Collinson (1988) suggested that the relatively small numbers of winged fruits or seeds suggested that few fruits and seeds blew onto the lake surface. Although the number of winged taxa represented in the collections has increased over the past few decades, all taxa still occur in low abundance (several just single specimens) further supporting this 11


suggestion. The distance of the source plant from the lake and transport-related processes of sorting due to specific weight or morphological characters are taphonomic filters that may have modified the plant macrofossil record in the oil shale. Rapid and differential decay of plant material under a paratropical climate may have contributed to a modification of the macrorecord in the oil shale (Wilde & Süss 2001) as may the presence or absence of resistant macromolecules in fruit walls and seed coats (Collinson 2011). In addition Wilde (1989, 2004) stressed the potential role of herbaceous communities which may have been developed at the margin of the lake. Vegetation with Cyclanthus, Volkeria, Araceae, Restionaceae and some ferns could have acted as a filter to impede entry into the lake of other plant remains arriving at the lake margin through overland runoff. Collinson (1988) and Wilde (1989) speculated that the forest probably approached the lake shoreline at least in some places and plant remains, including potentially many of the fruits and seeds, may have fallen directly into the lake.

Growth habits and vegetation reconstruction The picture of a tropical to subtropical (paratropical in the sense of Wolfe 1979) inland forest surrounding the early Middle Eocene lake at Messel which was suggested for the first time by Engelhardt (1922), has been confirmed especially by comparison to other nearby Middle Eocene sites (e.g. Wilde 1995, Wilde & Frankenhäuser 1998) and is now generally accepted (e.g. Wilde 1989, Schaal & Ziegler 1992, Gruber & Micklich 2007). At the time of deposition of the Middle Messel Formation the lake itself was a long-term (>600,000 years) stable meromictic lacustrine system with annually occurring blooms of Tetraedron (Goth 1990, Lenz et al. 2010). Collinson (1988) noted that there was no spatial localization in any of the fruits and seeds recorded in excavations at that time in sufficient numbers to enable consideration of their distribution at the various collecting sites. This wide distribution suggests that all of them occurred all around the lake. The taphocoenosis of the oil shale allowed Wilde (1989) to speculate about some kind of a zonation of the vegetation around the lake which is consistent with more recent data (Lenz et al. 2010). Inferences as to growth habit of the Messel plants are based on the habits of extant relatives. Although this requires the assumption of uniformitarianism, we believe it is a good approach for those taxa in which diverse extant representatives are consistent in their habit. For example, the extant genera of Menispermaceae and Vitaceae are dominantly lianas today. In addition, the tribes Phytocreneae and Iodeae of Icacinaceae, both well represented in the flora, are lianas. These same families and tribes are also well represented in the London Clay and Clarno 12

deposits, and assumed to indicate excellent representation of lianas in those Early and Middle Eocene floras as well. Although Rhamnaceae are variable in extant habit from shrubs to trees and lianas, the genus Berchemia Neck. ex DC. of the Messel assemblage, has species that are consistently climbers today. It is clear from the identified taxa that lianas are an important component of the Messel fruit and seed flora. Strikingly, at least 32 of the 89 species for which habit could be inferred are lianas (more than one third). This may in part reflect a bias for lianas to colonize light gaps and openings of the forest, particularly along stream and lake sides, where conditions for deposition and preservation are particularly favourable. The diversity and frequency of remains of families which are today dominated by lianas and climbing plants can nevertheless be interpreted in terms of a light-loving curtain covering the edges of the forest (both lake margins and clearings) and parts of its canopy (e.g. Wilde 1989). Other woody elements of the flora included a diversity of angiosperm shrubs and trees with a range of stature suggestive of a multiple canopy (rain?) forest. Trees likely included Ailanthus, Juglandaceae (3 genera), Lannea, Meliosma, Mytilaria Lecomte, Pleiogynium Engl., the mastixioids, some of the legumes, Nyssa L., Sloanea, and Tapiscia Oliv. Alangium, and Anacardium and others are variable in habit among different species today, ranging from shrub to tree or even liana. The frequently occurring leaves of Lauraceae may have been borne by understorey trees and shrubs. The presence of some ferns (e.g. Schizaeaceae) and monocotyledons such as Araceae (leaves only), Cyperaceae (fruits and pollen), Cyclanthaceae (fruits and seeds only) and Restionaceae (pollen only), suggests that herbaceous vegetation may have fringed the lake or existed in forest clearings. These herbaceous flowering plants may have flourished in an open marsh-like or swampy setting (Wilde 1989, Wilde & Micklich 2007). Angiosperm megafossils inferred to be derived from herbaceous plants are rare in most Eocene floras globally, and this may be attributed partly to taphonomic factors favoring the preservation of woody plants. Aquatic floating and submerged plants are apparently absent at Messel, with the possible exception of ?Nymphaeales. The single extant species of Decodon J. F. Gmel. is a woody plant typically rooted in aquatic areas. Wet ground or swampy conditions quite near the shoreline may have been preferred by the conifer Doliostrobus, the Pandanaceae, the palms and some other angiosperms (e.g. Nyssa, Ericaceae) (Collinson 1988) argued that palms grew close to the lake owing to their abundance at multiple collecting locations, the combination of occurrence of both delicate flowers and large fruits, and the persistent calyx on many fruits including specimens “in the process” of shedding their calyces. A few taxa of wind-pollinated plants are still only known from the pollen record, especially Pinaceae and Fagaceae. They presumably occupied more distant and probably


drier habitats (Wilde 1989; Lenz et al. 2011) that may even have been slightly more elevated.

Climatic interpretations Engelhardt (1922) already noted a tropical to subtropical character of the flora from the oil shale of Messel. Based on qualitative physiognomic considerations and the systematic composition of the leaves Wilde (1989) suggested a warm and humid, probably slightly seasonal but equable “oceanic” climate with mean annual temperatures between 25 and 30 °C for the Middle Eocene of Messel. This was recently supported by leaf margin analysis and a quantitative analysis (coexistence approach) of the systematic composition of the whole flora at family level by Grein et al. (2011) based on the family list of Wilde (2004). According to the coexistence approach, mean annual temperatures were between 16.8 and 23.9 °C with temperatures of the coldest month between 10.6 and 19.4 °C and temperatures of the warmest month between 24.7 and 27.9 °C. Mean annual precipitation is reconstructed to have been between 803 and 2540 mm and mean relative humidity between 73 and 77 %. Although we have not rerun the analysis, we think that it is unlikely that the newly recognised families herein (namely Alangiaceae, Altingiaceae, Burseraceae, Cannabaceae, Elaeocarpaceae, Myristicaceae, Rhamnaceae, Sabiaceae, Tapisciaceae and Toricelliaceae) will significantly alter the results derived from coexistence analysis. Thiele-Pfeiffer (1988) stated that she could not see obvious changes in the pollen and spore record along the 127 m of core Nr. 4 from 1980 that she studied. This statement must now be corrected in the light of a recent statistical analysis of her data (Lenz et al. 2005) and of the spores, pollen and resistant remains of algae from the more recent core Messel 2001 which clearly shows quantitative fluctuations in the range of Milankovitch cyclicity although no first or last appearances of taxa are observed (Lenz et al. 2011). They record distinct changes in the flora within and around the Middle Eocene lake resulting from distinct climate fluctuations over time ranges from less than 10,000 to about 100,000 years. Climate fluctuations in the range of ENSO cycles, shorter than10 years, have been suggested by analyzing the thickness of individual varve couplets (Lenz et al. 2011).

Biogeographic considerations Palaeobotany contributes greatly to our understanding of the changing patterns of plant distribution over time. In this section we consider the biogeographic implications of Messel taxa in two ways. First we compare with other major Eocene megafossil assemblages of the Northern

Hemisphere and consider their floristic similarities and differences to decipher biogeographic patterns and infer routes of biogeographic interchange. Second, we take the traditional approach of comparing the fossil assemblage with “nearest living relatives” with attention to the regions where those plants are native today. Comparison with other fossil floras The Messel flora provides an excellent vantage point for comparison with other mid-latitude Eocene floras around the Northern Hemisphere, because the range of plant remains preserved allows comparison with fruit and seed floras, like those of southern England and western North America, as well as with leaf floras like those of eastern Russia and China. Many of the families and genera represented in the Messel fruits and seeds are also common in the early Eocene London Clay flora (see summaries in Chandler 1964, Collinson 1983a, Collinson & Cleal 2001a) and in the Clarno Nut Beds flora (Manchester 1994).

North America Eocene megafossil floras of western North America (reviewed by DeVore & Pigg 2010, Pigg & DeVore 2010) are known mostly from lacustrine leaf assemblages. The fruit and seed flora of the Clarno Nut Beds (Scott 1954, Manchester 1994) provides the best known example of a western North American Eocene carpoflora for comparison with Messel. Manchester (1994) previously considered this flora to be mid-Middle Eocene based on fission track and 40Ar/39Ar dates of about 44 million years ago (mya) (Manchester 1994). However, Hanson (1996) considers the age of these beds to be early Middle Eocene, based on mammals from the Nut Beds site (which are interpreted to represent the Bridgerian land mammal age) and on an additional unpublished 40Ar/39Ar date by Swisher of 48.32 ± 0.11 mya (cited in Hanson 1996). Although the ages cited by Manchester and by Hanson for the Clarno Nut Beds are conflicting, they bracket the age of 47 my accepted for Messel. Fruit and seed genera shared between Messel and the Clarno Nut Beds include: Alangium, Anonaspermum (Ball) Reid & Chandler, Mastixia Blume, Pentoperculum Manchester, Bursericarpum Reid & Chandler, Mastixicarpum Chandler, Nyssa, Iodes? [cf. Palaeohosiea Kvaček & Bůžek], Palaeophytocrene, Pyrenacantha Hook. in Wight, Palaeocarya Saporta, Cruciptera, Laurocarpum, Leguminocarpon, Decodon(?), Magnolia L., Diploclisia Miers, Palaeosinomenium Chandler, Tinospora Miers, Tinomiscoidea Reid & Chandler, Meliosma Blume, Toricellia [“Tripartisemen” of Manchester 1994, revised in Manchester 1999], Tapiscia, Cleyera Adans., Aphananthe Link, Cedrelospermum, Parthe13


nocissus Planch., Vitis. Statistically, 19 % of the Clarno fruit and seed genera (28 of 145 genera) are shared with Messel, and 20 % of the Messel fruit and seed genera (28 of 140 genera; counting each Carpolithus entity as a distinct genus) treated here are shared with Clarno. These shared taxa apparently indicate continuity of populations across the North Atlantic Land Bridge (Tiffney 1985). Similarities between these floras are despite differences in depositional setting — lacustrine basin at Messel, vs fluvial/deltaic, with channel sands and slurry (matrix-supported conglomerates) in the Nut Beds deposit. Middle Eocene lacustrine floras of the Clarno Formation, for example that of the Cherry Creek drainage 20 km south of the Nut Beds site might be expected to share still more genera in common with Messel, given the lakebed depositional setting. The inventory of fruit and seed genera from the Cherry Creek flora includes 48 fruit and seed types (Manchester, unpublished observations at the Florida Museum of Natural History). Among these, Cruciptera, Cedrelospermum, Palaeocarya, Hooleya, Cypselites, and the tiny winged seeds that we call “Carpolithus 21” representing about 12 percent of the Cherry Creek lacustrine flora, are shared with the Messel flora. The rich leaf flora of the same beds shows very little overlap with Messel; Cedrelospermum, Ulmus L., and indet. Juglandaceae appear to be the only shared genera. Dominating elements of the Clarno lacustrine flora, including Betulaceae (Alnus Mill.) and two genera of Platanaceae (Platimeliphyllum N. Maslova and Macginitiea Wolfe & Wehr) are not known from Messel. Well-collected Eocene lacustrine floras of North America that can be compared with Messel include those of the Okanogan Highlands including the Republic flora of Washington (Wolfe & Wehr 1987, Wehr 1995, Wehr & Hopkins 1994, Wehr & Manchester 1996, Pigg & Wehr 2002), and McAbee flora of British Columbia (Dillhoff et al. 2005), and the more inland flora of the Parachute Creek Member of the Green River Formation of Colorado and Utah. These floras are best known from fossil leaves but also preserve occasional diagnostic fruits and seeds. The Republic and McAbee floras are rich in conifers (Schorn & Wehr 1996) and temperate angiosperm taxa including Fagus L., Alnus, Betula L., Acer L., Ulmus, Aesculus L., Fraxinus L., Dipteronia Oliv., Eucommia Oliv., Koelreuteria Medik. (See summary table in Dillhoff et al. 2005, DeVore & Pigg 2010). There is almost no overlap at the generic level with the Messel flora, except for rare Meliosma and Cruciptera fruits known from Republic, and relatively little overlap even with the geographically closer Clarno flora. Possibly the higher elevations inferred for these floras (Wolfe & Wehr 1987, Archibald & Greenwood 2005) may have favored frost-tolerant taxa that are not found in the Messel flora. The fruit and seed flora of the Princeton chert of South Central British Columbia (reviewed in DeVore & Pigg (2010) includes a few additional examples of taxa shared with Messel, including Lauraceae, Magnolia, 14

Lythraceae, Mastixiaceae, Vitaceae, and may yield other examples as research continues, but the abundance of Metasequoia and Pinus in the Princeton chert contrasts with the rarity of conifers at Messel. The well-collected early Middle Eocene flora of the Parachute Creek member of the Green River Formation in eastern Utah and adjacent western Colorado (MacGinitie 1969, Johnson & Plumb 1995) shares some genera with Messel, e.g. Cedrelospermum, Palaeophytocrene (only one specimen known, UF 15753–22924), Cypselites, Palaeocarya, but for the most part, this flora with its domination by Platanaceae (Macginitiea and Platanus L.), and Salicaceae (Populus spp., Pseudosalix Boucher, Manchester & Judd), is very distinct from that of Messel. Both Messel and the Parachute Creek flora, have leguminous pods and foliage, but they do not appear to represent the same entities.

Western Europe The London Clay flora includes at least 140 genera of fruits and seeds based on the compilation of Collinson & Cleal (2001a). It shares with Messel the genera Alangium, Steinhauera Presl, Pentoperculum, Lannea, Bursericarpum, Canarium Schumacher, Cyclanthus, Sloanea (Echinocarpus Blume), Euphorbiotheca, Corylopsis Siebold & Zucc., Icacinicarya Reid & Chandler, Iodes Blume (? cf Palaeohosiea), Palaeophytocrene, Laurocarpum, Leguminocarpum, Magnolia, Mimosites, Mastixia, Myristicacarpum, Diploclisia, Palaeosinomenium, Tinomiscoidea, Nyssa, Meliosma, Rutaspermum, Sapotispermum, Tapiscia, Vitis, Parthenocissus, Palaeovitis Reid & Chandler. Statistically, these 31 genera represent 22 % of the 140 Messel genera treated here (counting each Carpolithus entity as a distinct genus). It is interesting that the absolute number of genera shared between London Clay and Messel is about the same as the number of genera shared between Clarno and Messel. These similarities are despite different depositional settings between the marine deltaic deposition of London Clay versus inland lacustrine deposition of Messel and Clarno. Toricellia, which occurs both in the Messel oil shale and Clarno Nut Beds is absent from the London Clay flora. One possible explanation for this is the relatively small size which may mean specimens are overlooked during surface prospecting such as on the beach at Sheppey. However, this should not explain their absence in sieved residues of concentrates from Sheppey or from in situ clays e.g. Ockendon (Collinson 1983a). It seems possible that Toricellia was not growing within the catchment area of the London Clay deposits, perhaps because it was suited to more inland sites, as discussed in Collinson (1988: 192 under myrtalean seed cf. Lythraceae). Platanaceae fruiting heads are present in the Clarno Nut Beds but lacking in both Messel oil shale and the London Clay flora. In the London Clay the family is represented


by woods. This suggests that Platanaceae were not growing in the catchment area of the Messel lake. Annonaceae are not proven from the Messel oil shale but are diverse in the London Clay and Clarno Nut Beds floras. It is possible that (at least part of) the explanation for this lies in the different preservation states, with ruminate endosperm being most easily recognisable in permineralised internal casts that characterize the London Clay and Clarno fossils, vs externally preserved seeds in the Messel material. Based on experience collecting at Sheppey (MEC) and on NHMUK collections, two families, Salicaceae (Flacourtieae: Oncoba Forssk.) and Sapindaceae, are fairly common in the London Clay flora but are not known at Messel. Sapindaceae are characterized by distinctive morphologies of the fully developed pyritised embryos (Collinson 1983a) which would not be preserved in the Messel oil shale so hampering recognition of this family possibly for the same reasons that Annonaceae are difficult to recognise. Flacourtiaceae might not be recognisable from their external fruit morphology if compressed as would be the case in the Messel flora. Therefore these absences may be more apparent than real. Seeds with hairy appendages or awns and some winged taxa are recognised at Messel due to exceptional preservation (e.g. Ailanthus, Cedrelospermum, C. sp. 2, Cypselites, Cruciptera, Hooleya, and Palaeocarya) (see also Collinson et al. 2010). Their absence in floras such as the London Clay flora is likely to be more apparent than real. Genera with fruits diagnosed by wing morphology would not likely be recognised in the London Clay, because the mode of recovery (pyritized fruits liberated from clay by modern beach erosion) usually precludes wings. Such wings are preserved in the Clarno Nut Beds flora, however, so this factor might artificially increase the similarity of Messel to Clarno in relation to the geographically closer London Clay flora. The slightly younger taphocoenosis from Eckfeld in the Eifel Hills (Germany) (compilation in Lutz et al. 2010) has yielded a great number of plant taxa (Wilde & Frankenhäuser 1998). A number of fruits and seeds are directly comparable to taxa described from Messel (e.g. Pentoperculum, Cypselites, Canarium, Hooleya, Palaeocarya, Cocculus, Palaeosinomenium, Sloanea, Parthenocissus, Vitis). However, the material has not yet been studied in sufficient detail for closer comparisons. The Geiseltal area, near Halle an der Saale (Central Germany), which is famous for extremely well preserved Middle Eocene vertebrates, is another European site for fruits and seeds (Mai 1976), but the number of taxa described from sieved material is too low for statistical comparisons.

Eastern Asia Among Asian floras, the Fushun flora in Liaoning, northeastern China is perhaps the best known Eocene lacus-

trine flora (e.g. Wang et al. 2010a, b). Many leaves and some fruits of this flora were illustrated by Florin (1922), Endo (1942) and WGCPC (1978), although taxonomic determinations for angiosperms are much in need of revision. Although the Fushun oil shale preserves occasional remains of cones, fruits and seeds, it is best known for leaf remains. Therefore, when comparing these floras, it is important to take leaves into consideration as well as the fruits that are shared with Messel. Such comparison needs to include not only the previously published Messel leaf flora (Wilde 1989), but numerous additional undescribed fossil leaves in the Messel collections – a task beyond the scope of our current work. Radiometric dates from underlying strata in the Fushun Mine summarized by Wang et al. (2010a, b) indicate Paleocene age, so the leaf bearing strata might be Paleocene or early Eocene. Eocene age has been supported by palynological correlations, but the leaf bearing shales are not bracketed by radiometric dates; mammal and invertebrate fossils that would help to secure the age assignment are also lacking. Genera accepted by Manchester (unpublished observations from study of collections in Beijing and Nanjing) include Ailanthus, Alnus, Fagus, Acer, Deviacer Manchester, Craigia W. W. Sm. & W. E. Evans, Platimeliphyllum, Daphnogene Unger, Ulmus, Nordenskioldia Heer, Nyssidium Heer, Sabalites Saporta, Ginkgo L., Metasequoia Miki, Glyptostrobus Endl., Taxodium Rich., Cephalotaxus Siebold & Zucc. ex Endl., Pinus L., Salvinia Seg., and Lygodium Sw. Aside from a leaf of Cephalotaxus and fruits of Ailanthus, there is very little overlap between the Fushun flora and that of Messel, yet striking overlap with the Eocene floras of the Okanogan Highlands and Clarno Formation. The Fushun flora contrasts with the Messel flora by an abundance and diversity of conifers, and the common occurrence of deciduous Platanaceae and Betulaceae. The Huadian flora of Jilin Province, northeastern China, contains macrofossil leaves and occasional fruits in siltstones interbedded with oil shale (Manchester et al. 2005). Associated mammal fossils indicate a Middle Eocene age for this flora, which could mean that this flora is more age-appropriate for comparison with Messel than the Fushun flora. However, the Huadian flora shares little with Messel; Palaeosinomenium is the only convincing example from the fruit and seed flora. The concept of a band of Eocene “boreotropical forest” throughout the Northern Hemisphere (Wolfe 1977), has been widely accepted to explain the disjunct extant and fossil distribution patterns of thermophilic plant taxa. However, evidence for thermophilic vegetation in the Eocene of eastern Asia is weak (Collinson & Hooker 2003). The Eocene floras so far known from Asia appear to be cooler that those of Messel and London Clay. Perhaps the boreotropical forest was best developed in North America and Europe during the Eocene. The survival of components of this flora today in Asia might be a result of subsequent migration as the Turgai seaway receded. 15


Comparison with modern day floras The Messel species that we have been able to identify to modern genera show a range of modern distribution patterns. Some are elements of temperate deciduous forest today, like Magnolia, Tapiscia, Vitis and Parthenocissus, but a greater proportion show paratropical or tropical affinities, like Anacardium, Sloanea, and Phytocrene Wall. Geographically, the modern distribution patterns for genera identified from Messel fall into several main categories: eastern Asian, eastern North American, Old World tropics, Southeast Asia-Malesia, and Neotropics. Among examples extending into temperate climatic zones today, some of the Messel species represent species that are now distributed disjunctly around the Northern Hemisphere, e.g. Vitis, Parthenocissus and Nyssa. Decodon is restricted to eastern North America. Those of eastern Asian affinity include Diploclisia, Tapiscia, and Toricellia. Although Ailanthus ranges from India through Malesia to Australia today, the species most closely related to the Messel fossil is believed to be A. altissima (Mill.) Swingle of central China (Corbett & Manchester 2004). Alangium ranges from tropical Africa to Asia and Malesia today, but the species morphologically closest to the Messel species is A. chinensis (Lour.) Harms of eastern Asia today. It has been noted previously that a large number of European Tertiary genera have eastern Asian affinities today (Mai 1995, Martinetto 1998), and that many genera now considered endemic to eastern Asia had broader distribution in the Tertiary of North America and Europe (Manchester et al. 2009). In studying the early Eocene London Clay Flora, Reid & Chandler (1933) emphasized that, among taxa that could be identified to living genera, the highest proportion of closest relatives are found today in the IndoMalayan region. This is not the dominant pattern at Messel, but it is exemplified by Pleiogynium, with its modern species distributed from Malesia to the Pacific Islands. In addition, Messel genera with modern ranges extending from Eastern Asia to Malesia include Corylopsis, Mastixia, Mytilaria, and Phytocrene. Some representatives have broader tropical distributions today. Canarium, Cayratia Juss. and Stephania Loureiro are not found today in the New World but are widespread in the Old World tropics. Sloanea is widely distributed in the tropics today including Asia, Australia, Central and South America, and Madagascar. Palaeohosiea closely resembles Iodes which is an AfricanAsian disjunct. Lannea is distributed both in Africa and Indomalesia. Meliosma occurs in both Asian and New World tropics. Berchemia ranges from eastern Africa to eastern Asia, today but also occurs in western North America. A striking pattern of Neotropical affinities is illustrated by the genera Anacardium and Cyclanthus. Both of these genera are now confined to Central and South America in their modern distribution. The temporal and geographic 16

disjunction between the Messel representatives and the modern species in central and south America is intriguing because it indicates that these genera were once present in Europe but failed to survive in the Old World. It also suggests that these genera traversed the North Atlantic land bridge and formerly colonized North America and Greenland, although direct fossil evidence from those areas is lacking. A similar disjunction is evident in the mantid flies (Wedmann & Makarkin 2007), among the avian clade that includes hummingbirds and occurs at Messel (Mayr 2009), and among the phorusrhacid birds and horned toads (Schaal & Ziegler 1992). The mammal Eurotamandua joresi Storch, was originally interpreted as a representative of the neotropical xenarthran anteaters (Myrmecophagidae) but is now thought to be related to palaeanodonts in a basal position within the order Pholidota (pangolins) (Gaudin et al. 2009).

Future directions for Messel palaeobotanical research Various research projects are in progress on the Messel flora. S. Y. Smith is investigating Carpolithus species 57 and 58, to determine if their systematic affinity lies with the monocotyledons. MEC, and G. Storch are also continuing their work on the gut contents of Masillamys including Carpolithus sp. 57 in order to determine the diet of this rodent. S. Y. Smith and VW are investigating monocotyledonous leaves, including an assessment of whether any might have been produced by the plant bearing the Cyclanthus fruit. VW is investigating various specimens where fruits and seeds are attached to leafy shoots. Both of these studies will aid in the reconstruction of more complete Paleogene plants benefiting from the exceptional preservation at Messel. MEC and S. Y. Smith are evaluating the application of SRXTM to a wider variety of Messel fruits and seeds than is shown here. This work will determine if internal structural details can be obtained from rare specimens without destructive analysis. If so, this should help in the long term for recognition of the systematic affinity of at least some Carpolithus species. MEC is continuing her study of the fruits and seeds sieved from clastic sediments at site SMF 7 (Collinson 1988) most of which we hope can be identified readily in the light of this monograph. Studying the same taxon in both exceptional oil-shale preservation and more “typical” preservation from siliciclastics will expand the range of fossil floras which can readily be compared with that from Messel. MEC, VW and T. Borsch hope to bring their study of the Messel ?Nymphaealean flowers and their contained seeds to completion soon. However, the strong compression of the flowers is hampering our understanding of the floral organization and hindering our efforts to confirm the systematic affinity of these fascinating and common fossils, the largest flowers at Messel. In the long-


er term, we suggest that additional organic geochemical analyses should be carried out on selected Messel fossils to establish the former existence of biosynthetic pathways (e.g. Collinson & van Bergen 2004). Quantitative field censuses, recording every specimen at selected sites and stratigraphic levels, would be of value for further understanding of the spatial and temporal distribution of plants around the Messel lake. Such censuses could also record proportions of insect damaged leaves and seeds and might also provide evidence of repeated co-occurrence of leaves and fruits/seeds to guide the search for systematic affinities and enable more whole plant reconstructions. It would be very valuable to undertake field work focusing on very small plant remains (especially those smaller than 2 mm), as small fruits and seeds are probably biased against in the current Messel collections. A few taxa (e.g. Araliaceae, Symplocos) are absent from the fruit and seed flora but would be expected based on the occurrence of other organs. A targeted study of unidentified fruits and seeds might reveal these taxa. A reinvestigation of vertebrate gut contents in the light of the new information in this monograph should enable more dietary components to be identified, providing relevant seed anatomical features are evident in the specimens.

Systematics

Family Doliostrobaceae Kvaček Genus Doliostrobus Marion Doliostrobus taxiformis (Sternberg) Z. Kvaček emend. Kunzmann (Pl. 1a–c) Basionym: 1833 Cystoseirites taxiformis Sternberg, Flora der Vorwelt II, fac. 5 u. 6: 35, Taf. 18, fig. 1. Abridged synonymy: 1971 Doliostrobus taxiformis (Sternberg) Z. Kvaček, Palaeontogr. Abt. B, 135 (3–5): 118–123, pl. 31, figs 1–16, 19, pl. 32, figs 1–7; 1999 Kunzmann, Abh. Staatl. Mus. Mineral. Geol. Dresden, 45: 84–89, figs 15, 16, pl. 16–20.

Description: Individual cone scales obovate, lateral margins straight to concave. Maximum length 10–20 mm including distal tip (where observable); maximum width 10–15 mm, (reached in the distal third), narrowing to 2–3.5 mm at the basal attachment. Apical tip hook-like, length about 2–3 mm. Scale may appear striated due to surface topography caused by underlying resin canals, resin canals frequently exposed by fracture. Comments: A partial cone and leaves belonging to this species were described by Wilde (1989). Otto et al. 2002 argued that the terpenoid pattern of the resin supported a relationship to the “taxodioid” Cupressaceae s.l., especially Cunninghamia Schreb., rather than to

Araucariaceae. However, the ongoing debate on the systematic affinity of Doliostrobus (Bůžek et al.1968; Kvaček 2002a, b; Kunzmann 1999) is outside the scope of the present treatment. Seeds have not been found in the Messel material. Specimens: SM.B Me 20100–20102, plus those cited by Wilde (1989).

Family Alangiaceae DC. Genus Alangium Lam. Alangium sp. (Pl. 1d) Description: Endocarp bicarpellate, subovate in face view, rounded. Length 6 mm, dorsiventral width 4.3 mm. Surface smooth. Carpels subequal, the smaller (5 mm long), 5/6 the length of the larger. The two carpels separated by a longitudinal groove accommodating a row of several circular depressions (0.3–0.5 mm in diameter). Comments: The unequal development of two carpels and the row of circular depressions in a groove between the two carpels corresponds to the morphology of fruit stones in some species of extant and fossil Alangium sect. Marlea. The uniseriate row of circular depressions aligned with the septum corresponds precisely to the morphology seen in modern fruits of Alangium chinensis (Eyde et al. 1969: fig. 3f), and in anatomically preserved endocarps of Alangium eydei Manchester from the Middle Eocene Clarno Formation of Oregon (Manchester 1994: pl. 4, figs 5–13). The Messel species does not seem to show the strong surface striations of A. krutzschii Mai from the Geiseltal (Mai 1970, 1976), and is readily distinguished from A. dubium (Unger) Mai (1970) which has more than one row of circular depressions on the endocarp in the groove between the carpels. It is likely that this species is represented by additional specimens in the Messel collections, but they are difficult to recognise when compacted in face view; the specimen illustrated here was fortuitously compacted at right angles to the septum, facilitating recognition by the distinctive row of circular depressions. Specimen: SM.B Me 2215.

Family Altingiaceae Lindl. Genus Steinhauera Presl Steinhauera subglobosa Presl emend. Mai (Pl. 1e–f) 1838 Steinhauera subglobosa Presl in Sternberg, Versuch einer geolognostisch-botanischen Darstellung der Flora der Vorwelt, vol. 2, 7/8, 202, pl. 49, fig. 4, pl. 57, figs 1–4; 1968 Emended: Mai D.H., Palaeontographica Abt. B., 123: 190–196.

Description: Fruiting heads borne on a fertile branch with at least 7 flowering/fruiting heads. Infructescences com17


pact subglobose; peduncle short and thick (up to 3 mm thick). Heads up to 20 mm in diameter, composed of about 30 compartments (16 compartments visible on the exposed face). Compartments rounded-quadrangular in face view, with paired bracket shaped thickenings near the margin. Each of these units with two planes of symmetry, parallel and perpendicular to the bracket-shaped thickenings. These compartments, inferred to represent bilocular capsules, are embedded within a massive fibrous network with an essentially smooth surface. Capsules in face view rounded to quadrangular, 3–4 mm in diameter, intervening tissues 0.8–1 mm thick. There are no significant protrusions, indicating an absence of long styles. Comments: Globose infructescences of bilocular capsules characterize the extant genera of Altingiaceae, Altingia Noronha Liquidambar L., and Semiliquidambar H. T. Chang (Pigg et al. 2004; Ickert-Bond et al. 2005, 2007). Steinhauera Presl is a fossil genus accommodating isolated fruiting heads similar in construction to these extant genera (Kirchheimer 1943, Mai 1968), but distinguished from Liquidambar by the lack of persistent styles, and from Altingia which has both loculicidal and septicidal dehiscence (Chandler 1961a as Eoliquidambar; Mai & Walther 1985). Ickert-Bond et al. (2007) showed that Altingia infructescences are variously ornamented with short spines or tubercles in the regions between capsules. In contrast, the Messel specimens show these areas to be smooth and, in this respect, closely resemble extant Liquidambar orientalis Mill. (Ickert-Bond et al. 2005: fig. 5e). The Messel specimen shows for the first time the nature of attachment of the fruiting heads. They are borne helically on a stout woody axis, each head attached to the main axis by a short peduncle, although near the apex of the axis the immature heads may appear to be sessile (pl. 1e). Toward the apex of the reproductive twig the heads become progressively smaller (down to 9 mm in diameter), and perhaps less mature. It is not certain whether the distal heads are staminate (as in extant Liquidambar), or pistillate heads destined to become additional fruiting heads later in the same season. The surface morphology of the fruiting heads, with compartments having two bracket-shaped structures and no significant protrusions (pl. 1f), is particularly similar to that of specimens previously illustrated as Protoaltingia europaea Reid & Chandler (e.g. Collinson 1983a: fig. 99), from the London Clay and Eoliquidambar hordwellensis Chandler (1961a: pl. 26, figs 56–59) from the lower Headon Beds, which were placed in synonymy with Steinhauera subglobosa by Mai & Walther (1985). The type material of the species, from the Eocene of Staré Sedlo, was reillustrated photographically by Kvaček & Straková (1997), and seems to show more carpels per head than the specimens recovered here, but the extent of variability within the species remains to be documented. As part of this study MEC re-examined the London Clay specimen which she previously identified as Pro18

taltingia Reid & Chandler (collected and donated by Jim Craig, NHMUK V60822; figured in Collinson 1983a: fig. 99). The detail is indistinguishable from the Messel specimen detail on the plate [from SM.B Me 19093 (pl. 1f)]. [The next best London Clay specimen is V22419a where one or two surface compartments can be seen and support this. Other specimens are too fragmentary to comment]. Among other Messel fossils, the woody globose heads of this species are somewhat similar to those of the hamamelidaceous genus Mytilaria, known from Messel specimens with attached foliage (pls 12, 13). Whereas the Steinhauera heads are borne on branched axes (raceme like), the Mytilaria heads are borne solitary in the axils of leaves. There is also some similarity to Carpolithus sp. 36. Specimen: SM.B Me 19093.

Family Anacardiaceae R. Br. The Anacardiaceae are represented by at least four genera of fruits in the Messel flora: Anacardium, Lannea, Pentoperculum and Pleiogynium.

Genus Anacardium L. Anacardium germanicum Manchester, Wilde & Collinson (Pl. 2a–e) 2007 Anacardium germanicum Manchester, Wilde & Collinson, Int. J. Plant Sci., 168: 1199–1206.

Comments: Manchester et al. (2007) attributed this species to the genus Anacardium on the basis of the curved fruit and its inflated pedicel (hypocarp) (pl. 2a–d) which compare very favorably with the extant A. occidentale L. (pl. 2f). This is the only known European occurrence of the genus which is now native to Central and South America. Specimens: Holotype SM.B Me 7139. Others SM.B Me 2304, 7156, 7227 (hypocarp and lower part of fruit), 7279 (isolated fruit), 8737, 14414, 14755, 15048, 16021, 16846, 17041, 17770 (not 1770 as mistakenly cited in Manchester et al. 2007), 18095, 18993, 24045.

Genus Pentoperculum Manchester Pentoperculum minimus (Reid & Chandler) Manchester (Pl. 3a–h) Basionym: 1933 Dracontomelon minimus Reid & Chandler, London Clay Flora, 302, pl. 13, figs 20–24. Synonymy: 1994 Pentoperculum minimus (Reid & Chandler) Manchester, Palaeontogr. Amer., 36: pl. 5, figs 1–17.

Description: Fruit stones rounded-pentangular in transverse view, lenticular to obtrullate in lateral view,


5–7.2 mm equatorial diameter, ~3–7 mm high. Fruit consisting of 5 radially arranged single-seeded locules alternating with rounded lacunae. Each locule capped by an elliptical, longitudinally bipartite germination valve. Fruit surface usually smooth on the facets separated by ridges defining the pentagonal outline, sometimes with ten depressions visible on the lateral margins of the stone (pl. 3c, d) below the equator both aligned with, and alternating with, the five apical valves. Comments: The globose to oblate fruits of this species are found in different orientations in the shale, sometimes exposing the apical (pl. 3a, f), or basal (pl. 3b, g), or lateral (pl. 3c, d) surfaces. A transversely broken specimen clearly shows the radially arranged locules alternating with thick lacunate septa (pl. 3e). Laterally compacted specimens show the apical plugs and lateral depressions (pl. 3c, d). More detailed descriptions of this species, based on permineralized specimens, are provided by R eid & Chandler (1933 as Dracontomelon minimum) and Manchester (1994). The Messel specimens conform in size, pentangular morphology, position of elliptical apical germination valves, and the ten depressions on the lateral margins of the stone below the equator, to those known from the Lower Eocene of England. The same species is known from the London Clay (attributed to Dracontomelon in Reid & Chandler 1933), the Clarno Nut Beds (Manchester 1994), and from Eckfeld (Neuffer et al. 1996, Wilde & Frankenhäuser 1998). As noted by Reid & Chandler (1933), these fruits clearly belong to the Spondiadeae tribe of the Anacardiaceae, and are similar to those of extant Dracontomelon Blume. However, they are distinguished from that genus by the type of germination valves, which are solitary plugs in the modern genus, but bipartite shutter-like plugs in Pentoperculum (Manchester 1994). Similar bipartite shutter-like plugs occur in extant Pseudospondias Engl. and Haematostaphis Hook., but these genera have fewer locules and are longer than wide with prolate rather than oblate shape. Specimens: SM.B Me 2098, 4771, 5621, 8810, 11234, 21279, 21467.

Genus Pleiogynium Engl. Pleiogynium mitchellii sp. n. (Pl. 3i–m) Diagnosis: Fruit circular in polar view, c. 17 mm diameter, broadly lenticular in equatorial view, and about 6 mm high, although probably foreshortened through compaction (thickness in this plane suggests an originally oblate fruit being relatively tough and rigid). Fruit with c.10 pronounced longitudinal ribs. SRXTM reveals 10 locules, each aligned with these longitudinal ribs. Fully developed locules 1.2–1.5 mm wide, and 2.5–3 mm in dorsiventral

dimension, 3.5–4 mm high. On the apical face, there is a broad central depression (occupying almost a third of the diameter) from which the ridges radiate. Along each ridge is a central radially elongate elliptical protrusion, distinguished by difference in surface tissue. These areas are interpreted as opercula. Isodiametric cells occur on opercula surface in contrast to radially oriented fibers on either side of them. On the opposite face the fruit narrows uniformly toward a central circular attachment scar, 2.5 mm in diameter, with a central smaller ellipse, 0.4 mm diameter. Fruit surface between the ribs irregular. Sections through prominent vascular bundles are seen in digital sections arranged in a circle alternating with the locules. Etymology: This species is named in honour of John D. Mitchell for his research contributions on the morphology and systematics of extant Anacardiaceae (e.g. Mitchell et al. 2006). Comments: This fruit is much larger than the Pentoperculum specimens mentioned above and has twice as many carpels. Although only one specimen was observed, the application of SRXTM provided details of internal morphology (pl. 3l, m). Oblate fruits with multiple radiating locules occur in various families, including Euphorbiaceae (e.g. Hippomane L., Hura L.), Meliaceae, and Anacardiaceae. However, the presence of elliptical germination valves, or opercula, at the apical end of the fruit is diagnostic of spondoid Anacardiaceae (Mitchell et al. 2006). Among the extant genera of Spondiadeae, the number of locules, and corresponding germination valves, per fruit usually varies from one to five, but extends to 12 in Pleiogynium. This fossil corresponds especially well to the extant genus, Pleiogynium, which ranges from 5 to 12 carpels. Ten carpels is a common condition in P. timorense (DC.) Leenh. (pl. 3n, o). Specimen: Holotype SM.B Me 2227 (here designated, pl. 3i–m).

Genus Lannea A. Rich. Lannea hessenensis sp. n. (Pl. 2g–i) Diagnosis: Fruit usually elliptical to nearly circular in lateral outline, long axis 8.5–13 mm, short axis 7.2– 10.0 mm (n = 5). Surface smooth, outer fruit wall thin (0.1–0.2 mm). Outer endocarp surface covered with fine swirling fibres and basal attachment scar. Beneath this ornamentation are massive strands 0.3–1 mm wide composed of fibres which curve and branch enclosing 6–8 large deep smooth depressions (most frequent surface morphology encountered, outer layers having been lost). Curved germination slit frequently visible at apical end of fruit. Basal attachment scar circular, 2.4 mm wide, surrounded by a ring 0.2 mm thick; fruit faceted adjacent to the attachment. 19


Etymology: The species is named for the state of Hessen, Germany, where the Messel locality is situated. Comments: These fruits correspond to the anacardiaceous genus Lannea in the arrangement and orientation of prominent fibrous surficial strands and the overall swirling fibrous composition of the endocarp wall (pl. 2g–i). Some specimens show an oblique apical curved slit that appears to define the germination valve which is one of the characteristics of extant Lannea (pl. 2j). Although the number of locules and seeds was not directly observed, the fruits show only a single germination valve, indicative of a unilocular fruit. Lannea hessenensis closely resembles L. jenkinsii (Reid & Chandler) Chandler from the London Clay which is described as unilocular with woody gnarled endocarp, but, the L. jenkinsii fruits are smaller (8 mm long, 4.3 mm wide, 1.6 mm thick). The other London Clay species L. europaea (Reid & Chandler) Chandler, also reported from the Geiseltal (Mai 1976), differs by being two-loculed, with two operculae, and is still smaller, 6.6 mm long, 5.6 mm wide, 4 mm thick. Chesters (1957) described Lannea (as Odina miocenica Chesters) from the Miocene of Kenya. Those specimens are 8–10 mm long, 6–7 mm wide and 3 mm thick, one loculed by abortion and have fibres which curve and branch enclosing deep hollows, about 8 on each face. This material is similar to that from Messel but the fruits are narrower. More detailed comparisons are precluded due to the preservation of the Miocene specimens as mineralized casts. Specimens: Holotype SM.B Me 4882 (here designated, pl. 2h). Paratypes SM.B Me 2608, 4437, 4656, 4683, 4883, 5583, 7168, 7175, 7237, 7617, 7657, 7661, 8317, 8643, 8884, 15856, 17510, 19549, 21284, 21290, 21373, 21618.

Family Apocynaceae Juss. Genus Cypselites Heer Cypselites sp. (Pl. 4a–e) Description: Seed elongate-ellipsoidal with a terminal tuft of hairs. Seed body 5.7–10 mm long, 1.2–1.8 mm thick (n = 3). Seed body fusiform, narrowing to a pointed base, and rounded apically at junction with elongated tuft of hairs. Tuft of hairs at least two to three times the length of the seed body. Surface of seed body longitudinally striated with closely spaced rectangular cells in longitudinal rows. Comments: These narrow seeds, with a tuft of hairs arising from one end, are clearly adaptive for wind dispersal. As already noted by Reid & Chandler (1926), similar seeds occur in the extant apocynaceous genera Anodendron DC., Cleghornia Wight, Echites P. Browne, Forsteronia G. M ey ., Holarrhena R. B r ., Kibatalia G. Don, Odontadenia Benth., Pottsia Hook. & Arn., 20

Prestonia R. Br., Strophanthus DC., and Wrightia R. Br., but the distinction of these genera based only on seed characters is a difficult challenge. Among the extant Apocynaceae, comose seeds, with the hairs on the micropylar end, or the chalzal end, or sometimes on both ends of the seed, occur in all five subfamilies except the Rauvolfioideae (Endress 2000). The name Cypselites (Heer 1859) was published prior to, and has nomenclatural priority over Apocynospermum Reid & Chandler (1926). The name Echitonium Unger (1850a) was not only applied to such fossil seeds, but also to leaves of questionable affinity with the seeds. Cypselites neagelii is attributed to Heer 1859 (p. 2, pl. 101, fig. 1), and was based on a seed from the Miocene of Oeningen. Although the generic name might suggest affinity with Cypselis (Asteraceae), the tufted achenes of Asteraceae are readily distinguished from these tufted seeds by longitudinal ribs on the achene, and a more prominent articulation between the achene and the pappus. In the North American Tertiary, similar seeds occur in the Middle Eocene Green River Formation (Apocynospermum coloradensis Brown in MacGinitie 1969: pl. 18, fig. 4), and the Late Eocene of Florissant, Colorado (Manchester 2001). They are not known in North America after the Eocene, but in Europe they range from the Middle Eocene to the Miocene. In addition to the Messel locality, the seeds are found in the Late Eocene of England (Apocynospermum striatum Reid & Chandler, 1926), and Early Oligocene of Budapest-Obuda, Hungary (Hably et al. 2000, Botanical Department, Hungarian Natural History Museum Collection, BP 63.1039), and the České Středohoři Mountains (Kvaček & Walther 1995: pl. 9, fig. 2) the late Oligocene of Rott (Weyland 1938: 165, pl. 23, figs 11–13) and Aix (Saporta 1889: 53, pl. 8, figs 6, 17–19; pl. 20, fig. 6) and the Miocene of Switzerland and Randecker Maar (Rüffle 1963: 261). Occurrences were also reviewed in Collinson & van Bergen (2004). Specimens: SM.B Me 7624, 14650, 14661, 15469, 16422, 16901, 19824, 20180, 20301 (seed body lacking hairs). SM.B Me 4649 and 16422 (pl. 4d) probably belong to this taxon although the seed body reaches up to 17 mm long and 2.8 mm wide, the hairs are not distinct and the surface ornamentation is poorly preserved.

Family Arecaceae Bercht. & J. Presl Genus Friedemannia gen. n. Friedemannia messelensis gen. et sp. n. (Pl. 5a–n) Diagnosis: Fruit single-seeded, mature length 17.4– 25.8 mm, width 8.0–12.6 mm (n = 12), immature length 9.2 mm, width 4.0 mm (n = 1), frequently with persistent calyx, rarely seen in process of detachment of calyx, and commonly encountered lacking the calyx. Calyx of two whorls each of three segments, fibrous and longitudinally striate, covering one third to a quarter of the whole fruit, calyx length 6.1–7.7 mm, width 5.8–7.6 mm (n = 5).


Outer whorl of calyx one third to one half the length of the inner whirl. Fruit usually elliptical, sometimes ovate, with persistent short conical style (2–4 mm) protruding from a rounded apex. Outer fruit wall fibrous, in some cases fibres spreading as a result of degradation (SM.B Me 2123, pl. 5f). Seed structure was revealed by dissection of three specimens (pl. 5j–l). At the stylar end, large (3–4 mm) diameter circular hilar/chalazal plug, raphe extending from central protrusion down the full length of the seed to the attachment end. Fibrous bands (seen as grooves) radiate from the raphe. A single specimen (pl. 5m) reveals the hilar/chalazal plug through the partially degraded fruit wall. Etymology: The generic name honours the late Friedemann Schaarschmidt, who was first to recognise the palm affinities of these reproductive structures. Comments: These fruits are linked through an ontogenetic series including mature fruits (e.g. pl. 5d), young fruits (e.g. pl. 5n) to the flowers treated as Palmenblüten Typ A in Schaarschmidt & Wilde (1986). The fruits are found with and without the calyx. In the collections of mature well-preserved, fully prepared specimens showing relevant characters up to catalogue number SM.B Me 4999, there are 37 specimens with calyx and 78 specimens lacking the calyx. From field experience, these are the most abundant fruits in the Messel fruit and seed flora. Friedemannia fruits represented as much as 41 % of the propagules in the Senckenberg collections in 1988 (Collinson 1988). Remains of pinnate palm leaves of supposed lepidocaryoid affinity have been described from Messel (Schaarschmidt & Wilde 1986, Wilde 1989), but subsequently remains of a palmate leaf have also been found (unpublished).

convex, outer surface smooth. Texture inferred to have been leathery. Fruit apparently dehiscing in two valves, with two dense rows of subopposite seeds exposed (more than 8 per row, complete number unknown), seeds attached to a thickened axial structure (placental strand). Seeds borne on short stout funicles arising from the central placental strand. Placentation apparently axile. Inner surface of dehisced valve with striate margin (1.3– 2.1 mm broad) otherwise smooth; width of locular area 12 mm. Seeds obovate, smooth, with a distal elliptical structure (most probably chalaza) about 1.5–2 mm in diameter. No wings observed. Etymology: This species is named after the town of Darmstadt which is the county seat for Messel. Comments: The apparent axial placentation of these elongate fruits distinguishes them from the Messel legume pods. Moringaceae can have similar elongate woody valves, but the fruits of Moringa Adans. are trigonal, whereas these appear to have been bivalved. Although the seed outlines are clearly seen forming two longitudinal files within the fruit, we were unable to confirm the presence of wings on the seeds. The seeds may have been wingless, or the wings, which are typically membranous in this family, may not be evident because of preservational issues. Fruits of similar woody texture showing a similar pattern of placentation occur, for example, in extant Ceratophytum tetragonolobum (Jacq.) Sprague & Sandw. GH: Gentry & Morillo 10349, Venezuela (pl. 6d, f). Specimens: Holotype SM.B Me 7143 (here designated, pl. 6b, e); Paratype SM.B Me 7140.

Family Burseraceae Kunth 1824 Specimens: Holotype SM.B Me 2135 (here designated, pl. 5d). Specimens with calyx include SM.B Me 2123, 2125, 2126, 2130–2135, 2145, 2156, 2157, 2371, 2373–2375, 2377, 2378, 2380–2382, 2384, 2386, 2388, 2390–2395, 2397, 2399, 2400–2402, 2423, 2429, 2460, 2676, 2684, 4024, 4730, 4849; Specimens lacking calyx include SM.B Me 2348, 2367, 2403–2409, 2411–2417, 2419–2422, 2424–2426, 2431, 2432, 2435–2448, 2450, 2452–2454, 2456–2459, 2462, 2464– 2471, 2473, 2497, 2506, 2507, 2674, 2675, 2679, 2680, 2682, 4064, 4104, 4850–4853, 4857–4861, 4863, 4868; Dissected specimens and specimens with seed details SM.B Me 2137, 2138, 2147, 2148, 2157; immature fruit SM.B Me 2114. Miscellaneous additional specimens: SM.B Me 5193–5533, 5723–5725, 5747, 5748, 5753.

The Burseraceae are represented by at least two genera in the Messel Assemblage: Canarium and Bursericarpum.

Genus Canarium Stickman Canarium sp. (Pl. 7a–f)

Family Bignoniaceae Juss. Genus Darmstadtia gen. n. Darmstadtia biseriata gen. et sp. n. (Pl. 6a–c, e)

Description: Endocarp elliptical to oval in lateral view, length 12 mm, width across one of the flat faces 8 mm (n = 3). Prominently triangular in apical and basal view, with three prominent facets, flat to slightly concave, smooth. Each of the three keels with a very narrow groove. Base incomplete, apex of each facet protruding slightly. Germination valve triangular, apical.

Diagnosis: Fruit elongate ellipsoidal in outline, incomplete length of fruit 62 mm, width 15 mm. One end rounded, the other end broken, lateral margins smoothly

Comments: Fruits of this kind were initially assigned to the morphogenus Tricarpellites Bowerbank (e.g. Reid & Chandler 1933), but Gregor & Goth (1979) recognised 21


their affinities to Canarium. These fruits are extremely rare at Messel, but they correspond in all observable respects to the much larger population known from the upper Middle Eocene of Eckfeld for which anatomical study confirms affinity with Canarium (Wilde, Frankenhäuser & Mai, unpublished data). Canarium fruits are also found in the Early Eocene of Virginia (Tiffney 1999). Specimen SM.B Me 4610 (pl. 7f) shows the position from where the triangular germination valve has been shed. Specimens: SM.B Me 2300, 2329, 4610, HLMD-Me-15716.

Genus Bursericarpum Reid & Chandler Bursericarpum sp. (Pl. 7g, h) Description: Endocarp D-shaped in lateral view, with an almost straight ventral margin and an almost semicircular dorsal margin, 5.5–6.3 mm high, 4.6–5.0 mm dorsiventral thickness. Surface smooth. Placental slit almost transverse, extending from the ventral surface as much as half the distance towards the periphery. Immediately above the placental slit, a thickened wedge-like band extends a similar distance from the ventral margin, probably reflecting an underlying chalazal fibre band. Germination valve not observed. Comments: Bursericarpum species are known from the Lower and Middle Eocene of England and the Middle Eocene of North America (Chandler 1961a, c; Collinson 1983a; Manchester 1994). This morphogenus accommodates pyrenes of the kind found in several genera of Burseraceae today e.g. Bursera Jacq. ex L., Commiphora Jacq., and Protium Burm.f.

inflation of the basal portions of 4466 (pl. 7j) and 4476 (pl. 7l). These endocarps bear some similarity to those of Aphananthe, particularly in the subapical peg-like protrusion. They are smaller, however, than the extant species and smaller than the fossils named Aphananthe tenuicostata Dorofeev from the Oligocene of western Siberia (Dorofeev in Takhtajan & Zhilin 1982: pl. 110, fig. 6). The Siberian specimens are 4.7–5.7 mm long, whereas the Messel specimens are just 3–4 mm. Another difference is the angle of the peg-like protrusion in the Messel specimens it protrudes perpendicular to the body but in Dorofeev’s material it protrudes parallel to the surface (as in modern Aphananthe figured by Manchester 1989b: fig. 12.7 D). Not having carried out detailed anatomical study of this material, our assignment to Aphananthe remains tentative. Its affinities may be close to, if not within, this modern genus. The related celtoid genus Gironniera Gaudich. has also been reported in the palaeocarpological record, but those fossils do not display a prominent placental plug (Manchester 1989a). Fossil Moraceae and Cannabaceae have a terminal protrusion but this does not have a distinctive texture and is a funicular extension of the seed coat surface in Moroidea Chandler or a saddle-shaped plug in Cannabaceae (Collinson 1989b) and therefore differs from the Messel specimens. Specimens: SM.B Me 4169, 4466, 4476, 13049.

Family Cyclanthaceae Poit. ex A. Rich. Genus Cyclanthus Poit. ex A. Rich. Cyclanthus messelensis S. Y. Smith, Collinson & Rudall (Pl. 8a–e) 2008 Cyclanthus messelensis S. Y. Smith, Collinson & Rudall, Am. J. Bot. 95: 688–699, figs 1–10, 15–20.

Specimen: SM.B Me 18255, 23208.

Description: Endocarp small, ovoid to pyriform, length 3–4 mm, width 3 mm (n = 3), distorted (slightly crumpled and folded), thin-walled; with a prominent rounded, peglike, asymmetrically placed subapical protrusion of dense amorphous content [not crushed like the rest of the fruit]. Seed coat overlaps and thins at protrusion edge. Surface not shiny; texture minutely scabrate. Surface with weakly defined isodiametric polygonal cells up to 50 μm. Wall thickness c. 70 μm. Where visible in two specimens (pl. 7j, l), the wall is lined with a firmly attached thin, papery shiny cuticle.

Description: Discoidal fruiting cycles up to 6 cm in diameter with a central hole, radiating fibre strands, and a thickened outer rim. Some specimens have cuticle preserved. Fluorescence and LM preparations show a thin cuticle with square epidermal cells that become elongate towards the centre of the fruit. Stomata are scattered, paracytic, and do not occur in the inner region of the fruit cycle. Most specimens preserve numerous in situ seeds. These are basally attached to fibre strands, 2 mm in length with an ovoid seed body, elongate micropylar end, and short chalazal neck. Seeds observed under SEM show two cuticular envelopes. In situ seeds often have adpressed thickened bands that may represent remains of thickened anticlinal walls of an outer seed coat layer. Longitudinal ridges and transverse ribs form a hexagonal pattern on the seed. For more detailed description and discussion, see Smith et al. (2008).

Comments: The pyriform shape is inferred from the fold in the basal portion of SM.B Me 4169 (pl. 7i) and partial

Comments: Recognition of this genus in the European Tertiary has interesting biogeographic implications

Family Cannabaceae Martinov Genus Aphananthe Planch. Aphananthe cf. tenuicostata Dorofeev (Pl. 7i–l)

22


because extant Cyclanthus is distributed today in the neotropics (Smith et al. 2008). Several discoidal fruiting cycles with in situ seeds (pl. 8a–e), and occasional isolated seeds, have been found in the oil shale. These fossils are virtually identical in fruit structure and seed morphology to the now monotypic genus, Cyclanthus (Cyclanthaceae), although some taphonomic processes (such as compaction or pyrite growth) have altered the appearance of the external seed coat relative to that of extant C. bipartitus Poit. ex A. Rich. Analyses of Messel specimens in comparison with isolated seeds from the British Eocene led to the transfer of “Scirpus” lakensis, to Cyclanthus lakensis (Chandler) S. Y. Smith, Collinson & Rudall (2008). Specimens: SM.B Me 4879 a & b, 4880, 4881, 2004, 2005, 2006 a & b, 5742, 5743, 8203, 8204, 14007, 17929, 20028, 24734.

Family Cyperaceae Juss. Genus Volkeria Smith, Collinson, Simpson, Rudall, Marone & Stampanoni Volkeria messelense Smith, Collinson, Simpson, Rudall, Marone & Stampanoni (Pl. 8f–i) 2009 Volkeria messelense Smith, Collinson, Simpson, Rudall, Marone & Stampanoni, Am. J. Bot. 96: 1506–1518, figs 1, 4, 5a–c.

Description: Compact, elongate-ellipsoidal fruiting heads, length 3.9–12.0 mm, width 4.0–8.2 mm, with small, spirally arranged, imbricate fruits, fruits rounded ovoid to diamond-shaped in apical view. Eight to 35 fruits visible on exposed portions, implying a total of at least 16–70 in the complete fruiting head. Twelve specimens with evidence of peduncle; longest preserved peduncle (SM.B Me 16474) c. 13 mm long, appearing swollen for basal 2.5 mm. Isolated fruits 1.4–2.2 mm in maximum length and 1.1–2.2 mm in maximum width, obovoid with mucronate apex and truncate base pierced by a large central aperture. Fruits single seeded. Pericarp three-layered with an inner sclerotic endocarp, a thick parenchymatous mesocarp and a thin, often poorly preserved epicarp. Seed represented by a cuticle with basal placentation. No discernable structures (such as bracts) amongst or subtending the fruits. See Smith et al. (2009b) for formal description and detailed discussion. Comments: Isolated fruits and endocarps of this extinct genus of Cyperaceae are very frequent fossils in the European Paleogene (e.g. Chandler 1964; Mai & Walther 1978; Collinson 1983b; Collinson et al. 1993; Collinson & Cleal 2001a, b; Smith et al. 2009b) but the Messel specimens are the first examples with preservation of infructescences (Smith et al. 2009b). Specimens: Holotype SM.B Me 16474. Infructescences: SM.B Me 2099, 2100 a & b, 2101, 2102, 2104–2107, 2108 a & b, 2109, 2110, 2559, 2659,

4046, 4087, 4093, 4162 a & b (not 4163 a & b as cited in Smith et al. 2009b), 4208, 4236 (with a different plant fragment below it), 4642, 4698, 4759, 4998, 5075, 5081, 7197, 7200, 8212, 8266, 10001 a & b, 11164, 12292, 13084, 13877, 14748, 15578, 16255, 16570, 16675, 17129, 17982, 18059, 18091 a & b, 18092 a & b, 18123, 18177, 19802, 20192, 20826, 20977, 23188, 23215. Solitary fruits: SM.B Me 2208, 2219–2224, 2624, 2627, 2633, 4041, 4042, 4103, 4141, 4300–4308, 4310-4312, 4314-4342, 4344, 4386, 4454, 4457 (not 4497 as listed in Smith et al. 2009b), 4458, 4472, 4473, 4488, 4493, 4499, 4501, 4537, 4545, 4546, 4549, 4556, 4560, 4568, 4577, 4585, 4591, 4592, 4594, 4598, 4599, 4600, 4603–4605, 4608, 4609, 4621, 4623, 4629, 4672, 4720, 4735, 4758, 7691, 8093, 8101 (not 81011 as listed in Smith et al. 2009b), 8165, 8185, 8300, 8419, 8799, 11212, 12227 (incorrectly listed as 2227 in Smith et al. 2009b), 12278, 16067, 17344, 17982, 18235, 19006, 19557, 19674.

Family Elaeocarpaceae Juss. Genus Sloanea L. Sloanea messelensis sp. n. (Pl. 9a–o) Diagnosis: Fruit solitary, with a subtending disk, and long pedicel (19.5–63 mm length preserved (n = 3), about 2 mm thick). Mature fruit spherical to subglobose, without obvious apical protrusion. Height of fruit 8.5–23 mm (n = 5), width 9.5–24 mm (n = 6). Fruits ornamented with very fine closely spaced conical spinules. Fruit wall of two layers. Innermost layer (locular lining layer) is smooth and thin, normally less than 1 mm thick. Outer layer (2–2.5 mm; n = 2) comprised of closely packed anticlinally oriented spines, the tips of which form the surface ornament. When exposed in fractures, this layer always has a distinctively paler colour than other tissues in the fruits. Most specimens undehisced, without clear indications of dehiscence lines, however isolated valves (e.g. pl. 9i) and fruit portions (e.g. pl. 9a, h) with the same morphology and sculpture indicate longitudinal capsular dehiscence. Those valves have a median longitudinal septum indicating loculicidal dehiscence. Number of locules and carpels not immediately obvious, but radially oriented distal bulges could indicate five. Valve size in relation to fruit size suggests at least four or five valves. Seeds not known. Immature fruit retaining the style up to 4.46 mm long, elongate-triangular with broad base (1.15 mm wide narrowing to tip) and prominent perianth disk. Fruit in side view almost circular, 8.6 mm in diameter; large basal subtending disk covering about one quarter of the fruit and protruding at the margins; disk 9.1 mm wide and 3 mm high. Long peduncle preserved to 51 mm length, most of length c. 1 mm wide and broadening to 2 mm wide at the attachment disk. Comments: The anticlinally oriented, densely spinose outer fruit layer, prominent subtending disk and long pedicel correspond to those in modern and fossil Sloaneafruits (Kvaček et al. 2001, Manchester & Kvaček 2009). However, the Messel fossils differ from previously identified fossil representatives of this genus in having more densely packed spines. These fruits were treated as unde23


termined large fruits with striate outer layer by Collinson (1988). Etymology: The epithet refers to the Messel locality from which the specimens were collected. Specimens: Holotype SM.B Me 7133 (part and counterpart, here designated, pl. 9d). Paratypes SM.B Me 2049–2054, 4000, 4105, 4122, 4225, 4227, 4244, 4715, 7082, 7149, 7163, 7179, 7431, 8837, 13636 (immature with style), 14930,15466, 15475, 16420, 16542, 16443, 17450, 17451, 17521, 17775, 17797, 17898, 18076, 18161, 18947, 20039, 20053, 20449 (immature with style), 20537, 20607, 23881, 24312, 24318, 24331, 24406, 24425.

Family Euphorbiaceae Juss. Genus Euphorbiotheca Reid & Chandler Euphorbiotheca gothii sp. n. (Pls 10a–l, 11a–p) Diagnosis: Fruit spherical to oblate [in lateral compression breadth 5.8 to 9.9 mm, height 7.2–10.3 mm (n = 10)], 6.7–10 mm in diameter in polar compression. Fruits borne on a stout pedicel preserved to 5.9 mm long. Attachment area circular, without perianth remnants. Outer surface rugulate to verrucate at a submillimeter-scale, possibly leathery. Fruits radially symmetrical with six equally spaced meridional grooves. Meridional grooves delimit six equal segments which are frequently found shed, either singly or in pairs. Paired segments are interpreted as cocci, the units resulting from loculicidal dehiscence. A subsequent septicidal dehiscence (the initiation of which is frequently seen) divides the coccus into two equal single segments. The locule is positioned in the upper two thirds of the fruit. Locule outline obpyriform. Woody endocarp surrounds the locule, about 0.3 mm thick, and protrudes apically. Locule lining of the endocarp finely striate with striae radiating obliquely across the locule from the lower part of the fruit axis and arching apically toward endocarp margins. Endocarp surrounded by mesocarp tissue 1.5–2.2 mm thick with a finely wrinkled surface as seen on the sides of the capsule valves, suggestive of leathery texture, and a thin epicarp with verrucate to rugulate surface. Many of the fruits show evidence of ballistic dispersal, sometimes with cocci attached only by a thin portion of endocarp or pulled apart and/or twisted. Seeds flattened elliptical, 3.3 mm high, 2.2 mm wide, apparently one per locule. Etymology: This species is named after Kurt Goth, recognising his contributions to the study of Messel fruits and seeds. Comments: These tricarpellate fruits with both loculicidal and septicidal dehiscence are characteristic of Euphorbiaceae and segregate families (Komar 1992). The genus Euphorbiotheca is common in the London Clay flora, but none of those species are exactly like the Mes24

sel fossils. This taxon was indicated as the second most abundant fruit type in the Messel collections according to Collinson (1988). The capsules show evidence of explosive dehiscence like those of many extant Euphorbiaceae. The endocarp, as seen in profile view on the lateral faces of the cocci, has a T-shaped thickening apically along the ventral edge, with one arm directed a short distance apically and the other extending in the opposing direction (e.g. pl. 11e–g). This structural feature is seen in modern fruits of Euphorbiaceae and Phyllanthaceae with explosive dehiscence. Sometimes the cocci are twisted such that the apices of adjoining carpels are pointed in opposite directions (pl. 11g). Often the endocarp layer in isolated cocci is partially detached from the rest of the pericarp at the apical end (pl. 10g, j); sometimes the endocarp is completely missing from the pericarp and presumably detached during the dehiscence event [pl. 11c, f (left coccus)]. Detached endocarps are also preserved (pl. 11j). Most of the specimens lack seeds within the locules – not surprising given the explosive type of dehiscence indicated by this type of capsule. One specimen, however, shows the outlines made by a seed in an adjacent locule and the placentation being axile from near the base of the locule (pl. 11l), and another specimen clearly shows an intact seed remaining within the locule with the same placentation type (pl. 11n–p). The seed is elliptical and appears to lack a caruncle. Under high magnification a vascular strand (funicle?) was observed arching from near the apex of the fruit along the dorsal margin of the seed (but free from the seed) passing over the endocarp margin, then passing around the lower margin of the locule and rejoining near the axis. If it is correct that this species had just one seed per locule, we can rule out placement in the closely related family Phyllanthaceae (or Phyllanthoideae when treated as a subfamily of Euphorbiaceae) which contain two seeds per locule. The Euphorbiaceae sensu stricto, have just one seed per locule (Wurdack et al. 2005). In addition, the specimens that we have treated separately as Carpolithus sp. 7 (pl. 49g–j) could likely be immature fruits of this taxon. Although smaller and unopened, they are also on stout pedicels, have a similar surface texture and are divided by meridional grooves indicating six pyrenes in conformity with Euphorbiotheca gothii. They show strongly curved styles arising from the apex as in extant Euphorbiaceae. Although we only observed one (pl. 49g) or two styles (pl. 49h, i), we suspect that there were three, the third one being hidden within the sediment. The styles are basally connate and the stigmas are simple and unlobed. This organization is typical for Euphorbiaceae. Specimens: Holotype SM.B Me 7117 (here designated, pl. 10d). Paratypes 2007–2018, 2020–2038, 2040–2048, 2173, 2500, 2504, 2685–2690, 4002, 4009, 4010, 4017–4019, 4021, 4028, 4033, 4036, 4038, 4065, 4076 - has been sectioned but compaction has obscured internal details, 4085, 4088, 4094, 4131, 4135, 4137, 4139, 4112, 4155, 4173, 4200, 4214, 4247–4250, 4252–


4263, 4265–4289, 4291–4295, 4297–4299, 4550, 4567, 4584, 4597, 4606, 4630, 4700, 4705, 4717, 4718, 4783, 4787, 4799, 4807, 4938–4953, 5116– 5130, 5746, 5755 (?young fruit),7089, 7117, 7185, 7262,7267, 7268, 7269, 7283, 7285, 7358, 7360, 7362, 7309, 7380, 7386, 7581, 8063, 8076, 8264, 8297, 10560, 13838, 14908, 16493, 16561, 16684, 17044, 17327, 17492, 17511, 17552, 17779, 18054, 19479, 19537, 21531, 21672–83, 21685–87.

Family Hamamelidaceae R. Br. The Hamamelidaceae are represented by at least two fruit types, ie. Mytilaria and Corylopsis, and isolated seeds also referred to Corylopsis.

Genus Mytilaria Lecompte Mytilaria boglei sp. n. (Pl. 12, 13a–f) Diagnosis: Infructescences on thick peduncles axillary to palmately lobed entire-margined leaves with long petioles abruptly thickened at junction with lamina. Infructescence irregularly globular, composed of a stout central axis showing individual units in different developmental stages from top to bottom. Fully developed fruits are basal, but difficult to analyze because of preservation. Middle part of infructescence with numerous spirally arranged two-partite diamond-shaped pyramidal units diminishing in size distally [Lengths of immature diamond shape units 2.3 (n = 3) to 4.3 mm in mature part]. Each pyramidal unit has a transverse slit and a vertical groove and is embedded in a low but distinct smooth rim. Transverse slit of individual units perpendicular to the central axis of the infructescences. Apical part of central infructescence axis shows vertically elongate depressions, organized in the same spiral pattern as the bipartite units below but lacking bipartite structures inside. Styles are not obvious. Only a few of the gynoecia have enlarged into fruits although 12 diamond units preserved in face view in the upper part, indicating at least 24 potential fruits on the axes. Matured portions of the infructescences consist of enlarged woody dome-shaped units. In at least one case two such units are enclosed over their basal 1/3 by a thick cup-like structure with a flared rim.

fruit(s), and protruding on an elongate structure (peduncle) on either side of the main developed fruits. The undeveloped fruits form a regular pattern of diamondshaped domes as in the fossil. In palmate primary venation, palmate lobation with acute to rounded sinuses, and long petiole thickened at attachment with the lamina, the leaves attached with this fruit type are closely similar to the detached Messel leaves identified as Acer sp. in Wilde (1989). The two specimens available for that study were indicated as 5lobed, with three main lobes and two smaller ones. The leaves attached to the twig in pl. 13 vary from three to five lobed, and the five-lobed examples are architecturally similar to the one illustrated by Wilde (1989: pl. 18, fig. 5a). More detailed comparison will require analysis of the epidermal characters. Specimens: Holotype SM.B Me 18788 (here designated, pls 12a, 13 a, b, d–f). Paratype: SM.B Me 21325.

Genus Corylopsis Siebold & Zucc. Corylopsis maii sp. n. (Pl. 14a–e) Diagnosis: Infructescence spicate, consisting of a stout axis up to 1.5 mm thick, and up to 52 mm long, with helically arranged, sessile fruits. Individual fruit capsules rounded-quadrangular to subcircular in lateral outline (5–5.8 mm in height and breadth), opening loculicidally from the apex. Fruits with rough, irregularly verrucate surface, with a median longitudinal septal groove in a plane of bisymmetry extending to the apex. A pair of short, thick, persistent styles well separated from each other at the apex, with tips divergent and slightly outwardly curved. Cup-like structure (calyx) surrounding the base of each fruit, reaching about a quarter to a third of the distance from the base, with a smooth, unlobed margin. Number of fruits per axis up to 18 or more.

Etymology: This species name recognises the important contributions to Hamamelidaceae made by Lynn Bogle.

Etymology: This species is named in honour of Prof. Dieter Mai recognising his contributions to the palaeobotanical record of Hamamelidaceae, as well as his widereaching contributions to the Cretaceous and Cenozoic palaeobotany of Europe

Comments: A rare specimen showing leaves and fruits attached to the same twig proves the conspecificity of these head-like infructescences and alternate, palmately lobed leaves. The cuplike structure encircling the woody units is interpreted as hypanthium, as in other Hamamelidaceae. Extant Mytilaria matches these fossils in many respects (pl. 13g–m). The leaves are palmately lobed, entire-margined, with long, slender pedicels. The fruits are borne singly (or just a few) on a raceme of otherwise mostly undeveloped fruits. The positions of undeveloped fruits are seen over the lateral surface of the developed

Comments: Seeds missing from most specimens; a seed intact in SM.B Me 8954, but diagnostic features of the seed including hilar scar are partially hidden by fruit tissue. These bisymmetrical fruits, distinctive by their widely divergent paired styles (pl. 14e) correspond to capsules of the explosively opening type found in Hamamelidoideae. The fruits are not found isolated, but always in attachment with others along a thick axis. Usually the fruits have been compressed such that their plane of dehiscence lies parallel to the plane of the sedimentation. The external surface of each fruit is rough and 25


irregularly verrucate (pl. 14b, e) but sometimes the fruits have split along the dehiscence line, such that the fibrous construction of the wall and smooth lining of the pair of locules can be seen (pl. 14d). Endress (1989) listed 22 genera of this subfamily in a comparative study of flower and fruit structure of Hamamelidaceae. Since these fruits are sessile, they are assumed to develop from a spicate inflorescence of the type found in Tetrathyrium Benth., Neostrearia L. S. Sim., Noahdendron P. K. Endress, B. Hyland & Tracey, Ostrearia B aill ., Corylopsis, Sinowilsonia H emsl ., Fothergilla L., Matudaea Lundell, Molinadendron P. K. Endress, and Parrotia C. A. Mey. The hypanthium margin, i.e. the level at which the calyx is fused to the ovary, varies among different genera. In some genera it reaches to the apex of the fruit (resulting in inferior ovary): Sinowilsonia, Sycopsis Oliv., and Distyliopsis P. K. Endress. A few genera have superior ovaries without adnation of calyx (Distylium Siebold & Zucc., Matudaea, Parottia, Molinadendron). In most others the calyx reaches approximately half way (said to be half inferior ovary). In the Messel infructescences it extends only about 1/4 to 1/3 of the distance from the base to the apex (pl. 14c). The capsules in most genera of Hamamelidoideae split prominently in both the septicidal plane (separating the two seeds) and in the loculicidal plane, giving a four lobed appearance to dehisced fruits. In the Messel species, however, there is only minimal splitting in the septicidal plane. Among the extant genera we have compared to the Messel fossils, Corylopsis is the most similar. The extant species of Corylopsis that we examined had hypanthium fused to a higher level (about half the distance between base and apex of fruit), whereas the perianth of C. maii extends only about 1/4 of the distance, but our investigation included only a few extant species of this genus and it is likley that a broader survey of Corylopsis might reveal examples with shorter perianth in relation to fruit length. Hamamelis is also similar in fruit morphology, but the fruits of that genus do not aggregate in long spikes as seen in this fossil (Zhao & Li 2008). C. maii also resembles the fossil genus Hamawilsonia based on silicified infructescences with sessile, bihorned, squarish fruits from the Paleocene of North Dakota, USA (Benedict et al. 2008), but the Messel infructescences are only about half as large, and direct comparison is difficult due to the lack of exposed external surfaces in the permineralized specimens. Specimens: Holotype SM.B Me 14508 (here designated, pl. 14c). Paratypes SM.B Me 1894, 7142, 7315, 7434, 8954, 14383, 14718, 15472, 16263, 18552, 23743, 23961.

Corylopsis waltheri sp. n. (Pl. 14f–t). 26

Diagnosis: Seed ovoid, length 3.5–8.4 mm, width 1.5– 3.6 mm, with length/breadth ratio from 1.5–2.3 (n = 14). Seed with one end more or less pointed, sometimes appearing beaked and opposite end more rounded and often faceted. Seed slightly laterally compressed, with a smooth, slightly shiny surface. Surface sculpture microscopic pattern of polygonal cells (especially near hilum) to elongated cells with inflated outer periclinal walls. Elongate cells forming longitudinally striate surface pattern, cells ranging up to 0.2 mm long by 0.04 mm wide. More truncate end of the seed with a recessed hilar scar (0.6–1 mm wide, n = 4), trough-like and ligulate-ovate passing slightly over the end of seed, where it reduces in width to a narrow slit (0.1 mm wide), but not descending down opposite side. Scar extending for one third to one half of the length of the seed. A slight facet is seen on the face opposite the hilar scar but the seed coat surface is not modified over the facet. Etymology: This species is named in honour of Prof. Harald Walther recognising his contributions to study of the fossil record of Hamelidaceae and numerous other taxa in the European Tertiary. Comments: The shape and characteristic hilar scar indicate the family Hamamelidaceae (Hamamelidoideae). Three of the extant genera have large, very asymmetric hilar scars: Sinowilsonia, Ostrearia, and Corylopsis. Ostrearia australiana Baill. seeds are obovoid and the larger arc of the hilum is circular rather than elongate and narrow; Sinowilsonia henryi Hemsl. seeds are ellipsoidal and the larger arc of the asymmetrical hilum is circular. Seeds of recent Corylopsis (5 sp. investigated) are ellipsoid and have asymmetrical hilar scars with narrow arms. We refer the Messel seeds to Corylopsis based on their large, very asymmetrical, narrow hilar scar. Four of the extant species have very asymmetrical hilar scars confined almost completely to one side of the seed. The other side of the seed has a large, oblique, flattened or slightly depressed facet (Zhao & Li 2008). Co-occurrence, size and familial affinity suggest that some of these seeds may have been shed by the above-described fruits, however most of these seeds are too large to have fit in the locules of Corylopsis maii. This raises the question whether the discrepancy is due to differential shrinkage, or whether the fruits and seeds are actually representing different species (or even genera). Corylopsis venablesi Chandl. from the London Clay differs from this species in having a hilar scar extending onto both sides of the seed as seen in illustrations by Chandler (1961b), and Mai & Walther (1985). The same is illustrated for Corylopsis sp. from the London Clay figured by Collinson (1983a). The Messel seeds are similar in size and the hilum is comparable to an unnamed species (Hamamelidaceae gen.? of Reid & Chandler 1933) from the London Clay, but the latter specimen lacks details of the testa. Corylopsis waltheri


has a more strongly asymmetric hilar scar, and does not have an obvious hilar facet of the kind illustrated by Zhao & Li (2008) in extant species. However, many of our specimens are recognised by the presence of the hilar scar and the other face of the seed is still embedded in the shale and hence unseen. Corylopsis minima Mai & Walter 1985 and C.? latisperma Chandler 1961b have a hilar scar that descends down both sides of the seed. Corylopsis bognorensis Chandler 1961b cannot be compared because the hilar scar is not visible. Seeds of Hamamelidoideae are hard, smooth, spindle-shaped and are forcibly ejected from the capsules. Endress (1989) considered that the explosive dehiscence mechanism shared by all genera of this subfamily place constraints on the variability of fruit and seed shape, resulting in uniform morphology and making it difficult to distinguish fossil seeds at the generic level. However, there are differences in seed morphology among some of the extant genera (Grote 1989, Manchester 1994, Benedict et al. 2008). Grote (1989: 148–152) reviewed the seed morphology of 27 extant species in 16 genera of the subfamily: the seeds are ellipsoid or obovoid, and have an apical or subapical hilar scar which ranges from being symmetrical to very asymmetrical, and sometimes extends as arms down two opposite sides of the seed (up to 47 % or more of the seed length in Corylopsis). Specimens: Holotype SM.B Me 4722 (here designated, pl. 14f). Paratypes SM.B Me 2163, 2171, 2177, 2181, 2182, 4726, 4738, 4911, 5142–5147, 5149, 5151, 5630, 7127, 7129, 7130, 7709, 7922,7924, 7925, 7933, 7937, 7939, 7940, 7944, 20522 (specimens free from matrix).

Family Icacinaceae Miers Icacinaceae are well represented at Messel, with genera corresponding to the Iodeae and Phytocreneae tribes which today are restricted to the old world tropics. Collinson (1988) identified the presence of Palaeophytocrene and Natsiatum Buch.-Ham. ex Arn. (the latter reidentified as Palaeohosiea herein) at Messel. However she also identified some specimens as prunoid Rosaceae which are now recognised as Icacinaceae herein. Genus Palaeohosiea Kvaček & Bůžek Palaeohosiea bilinica (Ettingshausen) Kvaček & Bůžek (Pls 15a–k, m, n, 16a–c) Basionym: 1869 Amygdalus bilinica Ettingshausen, p. 55, pro parte, pl. 53, fig. 22 (non fig. 23) Synonymy: 1995 Palaeohosiea bilinica (Ettingshausen) Kvaček & Bůžek, Tertiary Research 15 (3): 125–126, pl. 1, fig. 15.

Description: Endocarps almost circular to elliptical in face view, length 9.5–15 mm, width 7–11.5 mm, with

rounded base and more pointed (asymmetrically pointed) apex, flattened in almost all cases without evidence of having been heavily compacted. A few specimens are cracked (e.g. SM.B Me 4919, pl. 15a). Asymmetrical apical protrusion inferred to be funicular, sometimes with a slit-like funicular scar. In a few specimens (SM.B Me 4026, 4327, 7341), a non-reticulate bulge protrudes upwards from the lateral face(s). Surface ornamented with a coarse reticulum of ridges. Some of the ridges more elongate and prominent, more or less longitudinal (typically three, sometimes four on the visible face). The reticulae are polygonal, some with free ending branches (typically 1 per field) often thickened at the end. Endocarp sometimes splitting in two halves along the plane of bilateral symmetry when the oil shale is split during collection (pl. 15d). Endocarps unilocular, locule wall sometimes appearing smooth and not papillate (SM.B Me 2357, pl. 15m, n), but papillae have been verified in SM.B Me 4652. Papillae 16–22 μm diameter. Underlying the papillate layer is a surface of smaller, more or less isodiametric polygonal cells 12–16 μm with somewhat undulatory walls. One specimen studied by SEM lacks preservation of papillae, which we attribute to degredation (pl. 15m, n). Comments: These endocarps show a reticulate surface sculpture characteristic of the Iodeae tribe of the Icacinaceae. The tendency of these endocarps to split along the plane of bisymmetry (e.g. SM.B Me 5625, pl. 15d) supports the placement in the Iodeae and distinguishes them from tribe Phytocreneae, also present at Messel. Similar reticulate sculpture occurs on endocarps of the extant genera Iodes, Hosiea Hemsl. & E. H. Wilson, and Natsiatum. In Hosiea, the mesocarp venation is free from the endocarp, so that the ridges are low and rounded compared to those of Iodes and Natsiatum (Kvaček & Bůžek 1995). Natsiatum has a smooth locule lining, whereas Iodes and Hosiea have a papillate locule lining readily observed by SEM and high magnification light microscopy (Kvaček & Bůžek 1995). Manchester (1994) reported that living Natsiatum herpeticum Buch.-Ham. has a papillate locule lining, but did not cite the individual herbarium specimen on which the observation was made, and we now consider this to have been based on an incorrectly determined specimen. More recently, he examined a second collection of N. herpeticum (A: Grierson & Long 3775, Bhutan), and observed smooth, nonpapillate locule conforming to the illustration of Kvaček & Bůžek (1995: pl. 2, fig. 7). In SM.B Me 4652 (pl. 16 a–c) little dark brown round shiny “pimples” occur both on the broken inner edge of the endocarp (descending into the specimen) and in an area where endocarp has fractured tangentially. The size and distribution of the pimples matches that of the papillae seen lining the locules of extant and fossil Iodes (Manchester 1994). Underlying these pimples is a golden thin layer of shiny equiaxial polygonal cells (lower right portion of 27


the image in pl. 16c). If the layer of pimples had been lost by decay, the underlying layer might have been misinterpreted as the locule lining and led us to conclude that the locule lining was smooth. Another specimen examined by SEM (pl. 15m, n) shows a smooth locule surface, but the absence of papillae might be due to degradation. This suite of specimens corresponds well in size and surface sculpture with Palaeohosiea bilinica from the late Eocene of England and Bohemia. Kvaček & Bůžek (1995) created the new combination Palaeohosiea bilinica. They placed Natsiatum eocenicum Chandler from Hordle, England, in synonymy with Amygdalus bilinica Ettingshausen from the late Eocene of Kučlín, Bohemia, under this new combination. This species is distinguished from two other species of Palaeohosiea by the common occurrence of blind ending ridges within the prominent slightly elongate and isodiametric meshes of endocarp ridges. They noted the presence of papillae (but “only rarely preserved”), 25 μm in diameter lining the locule. Although Kvaček & Bůžek clearly distinguished their new genus Palaeohosiea from extant Natsiatum and Hosiea, their comparison with extant Iodes seems to have been limited to a few extant species. They distinguished Iodes as having non-keeled endocarps with rounded oval profile and only low sculptural ridges. However we observed a well defined keel and prominent ridges in some species, e.g. I. seguini (H. Lév.) Rehder, so the hypothesis that Palaeohosiea and Iodes represent distinct genera needs to be revisited with a more complete survey of extant taxa. Similarly sculptured endocarps, also lacking papillae in the locules, have been described from silicified specimens in the Paleocene of North America as Icacinicarya dictyota Pigg, Manchester & DeVore (2008) and from impression specimens as Icacinicaryites linchensis Pigg, Manchester & DeVore (2008). These, too, will need to be reconsidered in the context of a broader survey of extant Icacinaceae. One of us (MEC) recently reexamined the British Paleogene material treated as Natsiatum eocenicum by Chandler (1961a, 1962), and confirmed that the endocarps do indeed have a papillate locule lining, as seen on V42152 from the type locality of Hordle and on V40618, V40621, V40622 from Lake. In V40618 the papillae were only visible in a small area of the well-protected locule e.g. in, slight infolds near the, margins, whereas where the locule lining revealed was essentially flat and exposed they seemed to be missing – possibly worn down by erosion of the exposed surface during fossilization or torn as the specimen split open and subsequently disintegrated. This observation indicates that the specimens from England may have closer affinity to extant Iodes, rather than Natsiatum, and conform to the fossil genus, Palaeohosiea as recognised by Kvaček & Bůžek (1995). Specimens: SM.B Me 631, 2079–2088, 2091, 2356, 2357 (studied by SEM for locule lining), 4007, 4026, 4073, 4156, 4327, 4390, 4391, 4393, 4394, 4595, 4616, 4619, 4652 (broken piece studied for locule lining), 4733, 4919–4925, 4927–4930, 4954–4987, 5625, 7320, 7327, 7341, 17596, 21584–21586.

28

Palaeohosiea sp. (Pl. 16d–j) Description: Endocarp subovoid, asymmetrical to symmetrical, base rounded to obtusely pointed, apex more angular than base, right-angle to obtusely pointed. Length 8.5–12.0 mm, width 5.0–8.0 (n = 3), thickness c. 2 mm (partly due to compaction), deduced to have been narrowly lenticular in cross section prior to compaction (thickness measured on SM.B Me 4626 when dried). Surface reticulate with a system of anastomosing ridges, enclosing polygonal areoles, some more or less isodiametric, c. 1 mm in diameter, others, particularly toward the apex, elongate longitudinally, up to 4 mm. Areoles sometimes with a single freely ending vein-ridge. Ridges usually with a narrow groove at the crest, and with aligned fibers. Wall less than 1 mm thick, homogeneous. Locule lining papillate. Papillae 18–24 μm in diameter. Comments: At first glance, these specimens seem to belong to the Messel Palaeohosiea bilinica (pl. 15). However, they lack the three or four stronger longitudinal ridges instead having an irregular reticulum all over their surfaces. They also possess some elongate fields in the reticulum, and lack the apical bulge marking the funicular canal. The apex is rather more symmetrical, and the base sometimes is slightly obtuse. The endocarp wall is homogeneous with a cellular pattern apparently aligned, not isodiametric. This is similar in morphology to Icacinicarya sp. 1 (see below), but much smaller. Endocarps of similar shape and sculpture occur in Icacinaceae and Rosaceae (Prunoideae), but the papillate locule lining indicates icacinaceous affinity. After photographing SM.B Me 4626 (pl. 16 e) in undamaged condition, we removed the specimen from glycerol, washed with water and dried it with heat from an incandescent lamp, forcing it to fracture thereby revealing the wall thickness and locule lining (pl. 16h). A small fragment removed for SEM shows the presence of papillae on the locule lining (pl. 16i). The papillae can also be seen by high resolution light microscopy. These are similar to those observed in a specimen we have attributed to P. bilinica (pl. 16, fig. c). Specimens: SM.B Me 2294–2296, 4089, 4388, 4389, 4392, 4626, 7112, 7323, 17702, 21587.

Genus Natsiatum Buch.-Ham. ex Arn. cf. Natsiatum sp. (Pl. 16k–m) Description: Endocarp ovate in face view, subrounded apically and basally, symmetrical, 10 mm long, 8 mm wide, thickness not known but inferred to be lenticular in cross section. Surface reticulate with a system of anastomosing ridges, enclosing more or less isodiametric po-


lygonal areoles, 0.8–2 mm wide, 1.1–3.8 mm long, sizes and shapes of areoles variable over the surface. Areoles containing numerous fine blindly ending branches from the ridges. Internal structure unknown. Comments: This endocarp morphotype differs from all other reticulately ridged endocarps of the Messel assemblage in having several fine thin blind ending branches in each field of the reticulum, a more symmetrical outline, and less dominantly longitudinal pattern of the ridges. In this feature of ramifying venation within the reticulae, this species conforms to extant Natsiatum (pl. 15l). Our hypothesis that this fossil represents Natsiatum predicts that the locule lining should be smooth, nonpapillate, but we have not yet investigated the internal structure of these two specimens. Following a comparative study of extant Hosiea, Iodes, and Natsiatum, Stull et al. (2011) recently recognized Natsiatum endocarps from the Eocene of Tennessee. Specimens: SM.B Me 4926, 17529.

Genus Icacinicarya Reid & Chandler Icacinicarya tiffneyi sp. n. (Pl. 17a–i) Diagnosis: Fruit obovate, 30–32 mm long, 19–24 mm wide, 5.5 mm thick, unilocular. Longitudinally ridged; with 7–10 main ridges per face, ridges from 2–7 mm wide. Ridge prominence diminishing towards the rounded, funicular end. Ridges anastomosing about halfway from the narrowed end, having fewer anastomoses and more elongate meshes at the opposite end; frequent anastomoses delimiting more isodiametric, polygonal meshes. Longitudinally aligned fibers occur only on the ridges. Other surface cells within the meshes black and shiny, some longitudinally elongate, others isodiametric, ca. 20–60 μm in length. Asymmetrical bulge with a horizontally elongate mouthlike scar near the apex. Fruit wall uniform, ca. 1 mm thick composed of dense “parenchymatous” tissue (not fibrous). Locule filled with tissue of isodiametric cells larger than those of the surrounding fruit wall. Etymology: This species is named in honour of Bruce H. Tiffney, recognizing his contributions in palaeobotany and education. Comments: This resembles the Icacinaceous species, Iodicarpa ampla Manchester from the Clarno Nut Beds (Manchester 1994) in the coarsely reticulate surface, thick wall, large size (average 31 × 26 mm) and asymmetrical bulge near the apex. The unilocular construction, confirmed by transverse sectioning (pl. 17 g) also corresponds. The asymmetrical apical bulge with transverse slit or scar (e.g. pl. 17a, b) is interpreted as the placental bulge where the funicle entered the locule, which is

similarly prominent in the extinct genus Comiclabium Manchester (1994). However, we have not determined whether the Messel specimens have papillate locule lining, interlocking sinuous walled cells in the endocarp, or a vascular bundle within the wall on one side in a plane of dehiscence. These characteristics are important to distinguish Iodicarpa and Comiclabium from Icacinicarya (Manchester 1994) and we therefore use the name Icacinicarya for the Messel fossils. Specimens: Holotype SM.B Me 5619 (here designated, pl. 17e–h). Paratypes SM.B Me 5557, 12911, 16650, 24055.

Tribe Phytocreneae Sleumer Genus Palaeophytocrene Reid & Chandler Palaeophytocrene sp. (Pl. 17j–p) Description: Endocarp wide elliptical in face view, nearly circular in transverse section; 7.5 mm high, 6 mm wide, 2.2 mm thick. Surface with small rather shallow, saucerlike pits/depressions (pl. 17k). Wall 0.6 mm thick (from digital sections), locule with stout protrusions of invaginated endocarp 0.4 mm long, 0.3 mm wide. A thin outer layer is dark brown and shiny with convex outer periclinal walls to the surface cells and this layer also descends into the pits (i.e. the pits are part of the overall topography). This outer layer has been removed (probably by preparation) over most of the exposed surface but present in patches and “sides” of the specimen. Underlying layer composed of cells which are isodiametric in surface view. Comments: This species has shallow, widely spaced depressions on the endocarp surface (pl. 17j, k). Digital sections, obtained by SRXTM analysis (pl. 17l–p), show that the specimen is unilocular with thick, short cylindrical protrusions of the endocarp into the locule. The outermost surface layer is black and shiny and has raised outer periclinal walls (looking like black pimples sometimes). This layer lines the depressions and is visible at the periphery of the specimen (pl. 17k) but has otherwise been lost over much of the “flattened” surface. The underlying layer has smaller cells equiaxial in surface view. Finely pitted endocarp surface and unilocular condition are among the characteristic features of fruits in the tribe Phytocreneae. Anatomically, the locular protrusions appear to be invaginations that are continuation of canals leading to the pores on the endocarp surface (pl. 17p). The surface topography of this species is very gentle compared to the deeply pitted sculpture of Pyrenacantha sp. 2, and the surface pits of this endocarp do not descend into narrow canallike structures such as we observed in that species (pls 18k, l; 19a–d). Based on relative wall thickness and proportions of the locule protrusions, this appears to correspond to the extinct genus Palaeophytocrene. For comparison, a transverse section of Palaeophytocrene pseudopersica Scott is 29


illustrated (pl. 19p). The locule protrusions are shorter and broader than those seen in extant Pyrenacantha (pl. 19m). By contrast, the endocarp pits of extant Phytocrene do not penetrate the locule (pl. 19n, o) Specimen: SM.B Me 21413.

Genus Phytocrene Wall. Phytocrene punctilinearis sp. n. (Pl. 18a, b, d, e, h–j) Diagnosis: Endocarp ellipsoidal, elliptical in face view, rounded basally, rounded to pointed apically, flat without traces of having been heavily compacted suggesting an original narrowly lenticular cross section. Length 19– 48 mm, width 12–30 mm. Surface smooth with 8–10 longitudinal rows of evenly spaced pits visible on exposed surface (suggesting a total of twice as many). Size of pits ranging from 0.5–1.5 mm in diameter. Each surface pit leading to a deep cylindrical invagination. Broken wall reveals that pits of the endocarp do not penetrate into the locule. Endocarp wall about 1.0 mm thick but only 0.3 mm in smaller specimens. Etymology: The epithet refers to the longitudinal arrangement of surface pits on these endocarps, one of the distinctions from Palaeophytocrene spp. Comments: Endocarps with pitted surface sculpture occur in the tribe Phytocreneae in Phytocrene, Pyrenacantha, and the extinct genus Palaeophytocrene R eid & C handler 1933, which is well represented in the Eocene of Europe and western North America (Rankin et al. 2008). Extant Phytocrene, in contrast with Palaeophytocrene, may have surface pits of the endocarp arranged in longitudinal rows (see Manchester 1994: pl. 19, fig. 16), and the tubercles or invaginations associated with these pits penetrate the endocarp wall without protruding into the locule (pl. 19o) or with only slight bulges into the locule (Sleumer 1971). Elongate, rod-like invaginations extend into the locule in Pyrenacantha (pl. 19l, m; Manchester 1994: pl. 19, fig. 15; Potgieter & van Wyk 1994), and shorter, thicker, peglike protrusions extend into the locule of Palaeophytocrene (pl. 19p; Manchester 1994; Rankin et al. 2008). The broken endocarp wall of SM.B Me 2326 (pl. 18b) does not show any protrusion of tubercles into the locules. This feature, combined with the longitudinal arrangement of pitting, leads us to place this species in Phytocrene (e.g. pl. 19n, o), a genus with about 11 species distributed today in SE Asia and Malesia (Sleumer 1971). The endocarp wall, up to 1 mm thick (pl. 18b) is thin in relation to the size of the endocarp, such that the fruits are preserved in a much flattened condition. SM.B Me 7150 (pl. 18e) is so thin-walled that the compressed thickness of the entire fruit is less than 1 mm. This contrasts with the more three30

dimensional preservation of other Messel Phytocreneae species placed in Palaeophytocrene and Pyrenacantha sp. 2. The only other known fossil occurrence of Phytocrene fruit is that of P. densipunctata Stull, Moore & Manchester (2011) from the Middle Eocene of Tennessee, USA, which has a similarly thin endocarp wall, but more densely spaced surface pits on the endocarp. If size is a meaningful specific character in this clade, at least two different species could be represented in the Messel material, the larger ones including SM.B Me 17899 (pl. 18a) and 2326 (pl. 18b) with a length of 44–48 mm, and smaller ones exemplified by SM.B Me 7150 (pl. 18e), 8449 and 12279 (pl. 18d) with a length of 19–19.5 mm. We consider it likely, however, that these represent specimens of one species in varying states of maturation. Specimens: Holotype SM.B Me 2326 (here designated, pl. 18b). Paratypes: SM.B Me 7150, 8449, 12279, 14047, 17899.

Genus Pyrenacantha Hook. in Wight Pyrenacantha sp. 1 (Pl. 18c, f, g) Description: Fruit drupe like with a soft tissue outer layer surrounding the endocarp. Endocarp pyriform, symmetrical to asymmetrical in apical portion, length 8.8–10.3 mm, width 6.8–7.6 mm, thickness unknown due to compaction. Endocarp surface dotted with more or less evenly spaced fine pits (0.2–0.3 mm in diameter), with a tendency to a pattern of even, radial, spacing of pits at the periphery (SM.B Me 4067, pl. 18f; 14490, pl. 18g). Endocarp thin-walled, such that the compressed fruit is less than 1 mm thick (SM.B Me 4067). Comments: Although the internal morphology of this species is unknown, the size and distribution of surface pits closely matches the condition of the extant species of Pyrenacantha illustrated photographically by Potgieter & van Wyk (1994). Only one specimen preserves the soft outer tissue of the fruit (pl. 18g), but it is significant in showing a lack of the prominent thick, stiff hairs characteristic of Phytocrene (Sleumer 1971), but rather a relatively smooth exocarp surface similar to that of extant Pyrenacantha species (Potgieter & van Wyk 1994). Pyrenacantha has “about 30 species, mainly African, but also extending towards Madagascar, India and the Philippines” (Potgieter & van Wyk 1994). Specimens: SM.B Me 4067, 14490, 16691.

Pyrenacantha sp. 2 (Pls 18k, l; 19a–d) Description: Endocarp wide-elliptical in face view, lensoidal in cross section, 12–14.5 mm long, 10–13 mm


wide, 7 mm thick (well inflated compared to Phytocrene specimens, but possibly foreshortened by compaction); rounded basally, obtusely pointed to rounded apically. Entire surface with more or less evenly and well spaced pits, not arranged in obvious longitudinal rows. Pits up to 1 mm in diameter, funneling to a canal 0.2–0.3 mm diameter (observed in SM.B Me 21431). Surface between pits smooth. About 10 pits across the width in face view. At one end there is a slight asymmetrical protrusion with a minute central cleft or pit giving rise to a circumferential groove. Displaced towards the margin of the opposite face is a deeply recessed elliptical scar bordered by a sharp margin which appears to be original (i.e. not damage and not a boring). An elongate grooved low ridge diverges from this depression and runs to the rounded end of the specimen. Comments: Like Phytocrene puctilinearis, described above, these endocarps are covered with numerous fine surface pits; however this species is not as greatly compacted, and thus was probably more robust and the pits in this species are not arranged in obvious rows. These pits are finer and more closely spaced than is typical of Palaeophytocrene. This endocarp closely resembles in surface morphology the endocarps of extant Pyrenacantha, for example P. grandiflora Baill. illustrated in Potgieter & van Wyk (1994). The latter species differs in the presence of a more pronounced basal scar, and we do not know the extent to which the tubercles protruded into the locule for this fossil. The fruits of this species are smaller than those of Pyrenacantha occidentalis Manchester from the Clarno Nut Beds of Oregon (Manchester 1994). The hypothesis that this species represents Pyrenacantha could be tested by sectioning to determine if the locule contained elongate protrusions of the endocarp as in the extant species and the Oregon fossil (Manchester 1994, Potgieter & van Wyk 1994). This species is distinguished from Pyrenacantha sp. 1, described above, by its large size, greater inflation, and more blunt apex. Specimens: SM.B Me 4992, 8464, 21431.

cf. Pyrenacantha (Pl. 19e) Description: Endocarp ellipsoidal, elliptical in face view, 22.5 mm high, 18 mm wide, rounded on both ends, c. 4–5 mm thick in the compressed state of the fossil, inferred to have been lenticular in cross section. Surface with numerous evenly spaced pits (about 25–30 pits across width at largest diameter). Pits approximately 0.2–0.3 mm in diameter. Comments: This fruit retains three dimensionality, contrasting with the flattened fruits of Phytocrene punctilinearis, and Pyrenacantha sp. 1. This suggests a relatively thick-walled fruit as in Palaeophytocrene, but

the spacing of surface pits is more dense than is known for previously recognised species of Palaeophytocrene. This species is similar in endocarp shape and size, and in the spacing of surface pits, to the endocarps of extant Pyrenacantha kaurabassana Baill. (Potgieter & van Wyk 1994: fig. 1f). It is distinguished from Pyrenacantha sp. 2, described above, by larger size and greater number of surface pits. The depth of protrusions into the locule is unknown for this fossil, so the placement in Pyrenacantha is provisional. Specimen: SM.B Me 7165.

Genus Icacinicarya Reid & Chandler Icacinicarya densipunctata sp. n. (Pl. 19f–k) Diagnosis: Endocarp large, wide-elliptical to nearly circular in face view, lenticular in transverse view, 27– 35 mm high, 27–28 mm wide, and 8 mm thick, lenticular in cross section. Small apical protrusion may represent a funicle. Endocarp not greatly compacted and very robust, indicating a strong woody endocarp tissue. Internal structure and wall anatomy not observed. Surface covered with fine closely spaced, randomly arranged pits of variable size, all less than 500 μm in diameter. Etymology: The specific epithet refers to the closely spaced surface pits. Comment: This species is distinguished by large size, combined with very small and closely spaced surface pits. Relative to endocarp size, the surface pits are smaller and more densely spaced than in previously known species of Pyrenacantha, Phytocrene and Palaeophytocrene. The apical bulge (presumably funicular) shown nicely in SM.B Me 23388 (pl. 19f, g) is not obvious in 14938 (pl. 19i, j), but both specimens show identical surface pitting (pl. 19h, k). Reid & Chandler (1933) intended the genus Icacinicarya for fossil species possessing endocarps and seeds with characters of the Icacinaceae that cannot be placed definitely to a particular modern genus. Until more information becomes available concerning the internal structure and anatomy of this species, it seems reasonable to place it within Icacinicarya. Specimens: Holotype SM.B Me 23388 (here designated, pl. 19f–h); Paratype SM.B Me 14938.

Icacinicarya sp. (Pl. 20a–c) Description: Fruit/endocarp circular in face view (diameter 14.5 mm), thickness in plane of compaction indicates 31


originally more or less globose. Entire surface ornamented with somewhat irregularly spaced flat-topped conical tubercles approximately 0.5 mm wide, up to 1 mm apart. Each tubercle has a central shallow straight sided craterlike pit. Surface between tubercles granular with very fine cellular pattern. No obvious style or attachment scar can be distinguished. Comments: The endocarp shape is similar to that of Pyrenacantha sp. 2 but the tubercules protrude, rather than being recessed as pits. In this respect, the morphology of the tubercles is unlike any other tubercled taxon at Messel, but it could possibly represent a different state of preservation of the same P. sp. 2. The tubercles might represent bases of abscised spines, or they may have had a glandular or secretory function. Further study of this single specimen by digital sectioning would potentially provide useful information about the tubercle structure. Specimen: SM.B Me 10564.

Family Juglandaceae DC. The walnut family is well represented in the Messel assemblage, represented by four types of foliage (Wilde 1989), more than a dozen pollen species (Thiele-Pfeiffer 1988), at least three fruit genera – Palaeocarya, Cruciptera, and Hooleya, and one superficially similar fruit – Carpolithus sp. 54. Engelhardt (1922) identified three Carya species from Messel as nuts: C. costata Ung. (pl. 32, fig. 12), C. ventricosa Stbg (Engelhardt 1922, pl. 34, figs 13–15), and a new species that he named Carya hickoryaeformis (Engelhardt 1922, pl. 32, fig. l). We have reexamined his material but were not able to confirm their identity as Carya Nutt. Nevertheless, Carya is recognisable in the Messel assemblage from pollen (Caryapollenites triangularis and C. circulus; Thiele-Pheiffer 1988).

Genus Cruciptera Manchester Cruciptera schaarschmidtii Manchester, Collinson & Goth (Pl. 20d–g, Pl. 21b) 1994 Cruciptera schaarschmidtii Manchester, Collinson & Goth, Int. J. Plant Sci., 155: 389, figs 1–10.

Description: Winged fruits with globose central nut 3– 6 mm diameter and four orthogonally arrayed, strap-like bract-derived wings arranged in a plane perpendicular to the long axis of the nut. Wings 2.7–4.5 mm wide, 8.5– 13 mm long. Maximum length of fruit measured from wing tip to wing tip, 28 mm. Four epigynous tepals arising apically from the nut. Wing venation open dichotomous consisting of a single order of closely spaced subparallel veins. Pedicel 1.8 mm long. Fruit and wings covered with circular peltate scales (Manchester et al. 1994). 32

Comments: This species has a fruit form convergent with that of the malpighiaceous genus Tetrapteris Cav., and was initially identified as “Tetrapteris sp.” (Collinson 1988: 189). Nutlet morphology, particularly well preserved in uncompressed specimens from the Clarno Formation of Oregon (Manchester 1991), and peltate scales well seen in the Messel specimens by epifluorescence on the surface of the nut and wings (Manchester et al. 1994), as well as the four epigynous tepals, and wing venation similar to that of extant Pterocarya, confirm the position of this taxon within Juglandaceae. Although Cruciptera has winged nuts rather similar to extant Pterocarya, it possesses four, rather than two, distinct wings. The wings are oriented perpendicular to the long axis of the fruit (rather than oblique as in Pterocarya), as is the single disk like wing of extant Cyclocarya. C. schaarschmidtii was fully discussed by Manchester et al. (1994), but many additional specimens have come to light since then, including the cluster of closely associated fruits (pl. 21b). Specimens: Holotype SM.B Me 7152. Others: SM.B Me 1893, 1948, 2060, 2061, 4110, 4229, 4240, 4876–4878, 4989, 7141, 7223, ?7239, 8381, 13819, 15449–15451, 15454 (lateral compression), 15471, 15528, 15552, 16266–16270, 16423, 16605, 16884, 17520, 18132, 18203, 18229, 19127, 19757, 19817, 19818, 19820, 20030, 20031, 20048, 20099, 20750, 21264, 21621, 21623, 24342, 24511 (7 fruits associated with a twig).

Genus Hooleya Reid & Chandler Hooleya sp. (Pl. 21a) Description: Fruit an ovoid nut, 4 mm high, 3 mm wide, with two large rounded lateral wings, 4.3 mm wide, 7.5 mm high, situated in the same plane as the nutlet-axis. Venation fan-like, bifurcating about half way, anastomosing near the margin. Nut and wings covered in numerous simple hairs and scattered peltate scales. Comments: There is still only a single specimen of this taxon in the Messel collections. Frankenhäuser & Wilde (1994) described numerous similar fruits from the slightly younger Middle Eocene of Eckfeld. The large assemblage of well preserved specimens demonstrated that the stylar region is very variable but compaction made it difficult to fully understand this feature which is critical for specific determination and systematic assignment to either Hooleya or the pterocaryoid alliance. Unfortunately the single Messel specimen does not have the stylar region preserved, but all other characters fall within the range of variation of the Eckfeld material. Following Manchester et al. 1994 we use the name Hooleya sp. Hooleya also occurs in the late Eocene of Sotzka (Unger 1850b), the latest Eocene Insect Limestone flora in England (Reid & Chandler 1926, Collinson et al. 2010), and the Late Eocene of Kučlín, Bohemia (Kvaček 2002c).


Specimen: SM.B Me 2064.

Genus Palaeocarya Saporta “Palaeocarya sp.” sensu Manchester et al. (Pl. 20h, i) Description: Fruits with prominently trilobed wings and a fourth less conspicuous (not always preserved) shorter and broader wing covering the other face of the fruit. Wing lobes with three three main veins including midvein and a pair of strongly ascending marginal veins. Comments: Following Manchester et al. (1994), these fossils are assignable to Palaeocarya Saporta, to the species P. macroptera (Brongniart) Jähnichen, Friedrich & Takáč 1984, but this taxon is obviously heterogeneous (Dilcher & Wilde, unpublished data) and therefore we refrain from making a formal species assignment. Contrary to the implication of the name, Palaeocarya Saporta is unrelated to the extant genus Carya; rather this is the fossil generic name having priority to accommodate winged nuts with the wing formed from a trilobed primary inflorescence bract, as in the extant genera Engelhardia Lesch. ex Blume, Alfaropsis Iljinsk., and Oreomunnea Oerst. The first of these genera is distributed in Asia and Malesia today, while the second and third are confined to tropical America. Some species of Palaeocarya combine bract venation characteristics of Oreomunnea with nutlet and style characters of Engelhardia.

(SM.B Me 5112, pl. 22g) and show cuticle thickness but do not resolve detail of other pericarp tissues. Comments: The large attachment scar and preservation of thick, tough, wrinkly cuticle, and oil cells indicate Lauraceous affinity. One of these was figured as Podocarpus sp. by Engelhardt (1922; HLMD-Me-1835, pl. 22j). It is not possible to compare with the majority of previously described species of Laurocarpum which are mostly based on permineralized material (e.g. Chandler 1964, Manchester 1994). Mai & Walther (1985) recognised three unnamed species of Laurocarpum from compressed fruits from the Weißelster Basin. This species is similar to their spec. 1 in overall morphology and size. Specimens: with scars visible: HLMD-Me-183, SM.B Me 2310 (pl. 22e, 18 mm, scar 15 mm), 2561 (pl. 22h, 12 mm wide, scar 7 mm wide), 4001 (side view), 4993 (pl. 22d, 16 mm, scar 11.5 mm in diameter), 4696 (15 mm wide, oblique side view, scar 12 mm), 4997 (pl. 22k, oblique side view, 12 mm wide, ca.10 mm high, scar 8 mm wide), 5102 (pl. 22i), 5112, 5113 (pl. 22l), 7153 (oblique side view, 17 mm wide, 11 mm high), 7317, 8386, 8396, 8411, 13000 (pl. 22b), 16235, 21616.

The following specimens show a cuticle exactly like this taxon but the position of their scar is embedded in oil shale. A few specimens of this morphology (included in specimen list above) were removed from oil shale and scar was proven to be present so it is likely (but not proven) that all these specimens belong to this taxon. SM.B Me 460, 2562, 2563, 4994, 5103, 5104, 5106, 5107 (very large example), 5108, 7599, 7377, 7411 (large example), 18088, 21617.

Specimens: SM.B Me 2065, 4230, 4239, 4712, 4988, 4990, 5599, 8382, 8402, 8407, 8694, 8912, 10623, 16265, 17402, 17483, 20032, 20683, 21371.

Laurocarpum sp. 2 (Pl. 22n)

Family Lauraceae Juss.

Description: As for Laurocarpum sp. 1 but about half the size, e.g. 6.9 × 6.3 mm (SM.B Me 4416, pl. 22n) with proportionately smaller scar.

We recognise at least three representatives of Lauraceae among the fruit collections, but more may reside among the globose nondescript specimens that have not been investigated in detail. Wilde (1989) accepted more than ten species of Lauraceae based on foliage.

Comments: There seems to be a clear break in the sizes of these two morphotypes (Laurocarpum sp. 1, 2) but otherwise they are too similar to distinguish. They may be two species of the same genus, or represent different stages in ontogeny (Little et al. 2009).

Genus Laurocarpum Reid & Chandler Laurocarpum sp. 1 (Pl. 22a–m)

Specimens: SM.B Me 2560, 2568, 4416 and possibly 4835, 5097, 7755 (pedicellate).

Description: Fruit originally spherical, slightly wider than high, from 12–18 mm in maximum dimension, with a large scar at the assumed base. Scar 7–15 mm in diameter (n = 5), perfectly circular, smooth edged, surface of fruit with thick smooth but strongly wrinkled cuticle. Some specimens (SM.B Me 7377, pl. 22a) show tiny regularly spaced yellow (can range white to red) dimples on the outer surface interpreted as oil cell contents. Digital sections (micro CT) indicate presence of a single locule

Laurocarpum sp. 3 (Pl. 21e–g) Description: Fruit circular in outline, 17 mm in diameter, with large central cavity 12 mm in diameter, and thick pericarp, 2.5–3 mm thick. Outer surface with a black, cuticular layer. Pericarp including an inner uniseriate layer of well lignified columnar cells 110–125 μm thick, probably an endocarp. Most of the pericarp tissue represented 33


by a thick layer of small isodiametric cells (smaller than those of endosperm/cotyledon) with interspersed darker star-shaped sclereids, interpreted here as mesocarp. Comments: The specimen has been broken open (uncertain whether transversely, longitudinally, or obliquely) showing circular outline, thick pericarp, and single circular locule. The inclusion of stellate-organised cells within the mesocarp, along with the internal columnar layer (pl. 21g), are diagnostic features of Lauraceae (Reid & Chandler 1933). Laurocarpum sp. 3 is distinguished from Laurocarpum species 1 and 2 by having a black outer cuticle, pericarp tissue differentiation and columnar cell layer (probable endocarp).

margins parallel without constrictions, tapering abruptly at the apex and base. Base slightly asymmetric, acute to rounded, apex rounded. Both margins slightly thickened with the one on the placental side slightly thicker than on the other. Seed chambers 7–18, rectangular (6–8 mm × 14–16 mm), elongate across width of pod, well delimited by transverse ridges about 1 mm thick. Seed outline ovate. Seed outlines are filling the width of the chambers, but usually extending only approximately 2/3 the distance towards the non-placental margin, seed outlines sometimes entirely filling the chamber (SM.B Me 19098). Venation seen only in a very small part of SM.B Me 4884, coarse veins reach to centre of pod from the placental margin, branch singly and obliquely into short lateral veins of almost equal stature. A fine reticulum is evident over the centre of the pod.


Lauraceae genus indet. 1 (Pl. 21c, d) Description: Fruit globose, 4 mm in diameter, with an abruptly acute apex. Flattened by compression, preserved with a smooth dark cuticle covering through which can be seen closely spaced yellow dots. Comments: The cuticular preservation is typical for Lauraceae. The evenly spaced yellow dots are interpreted to represent resin from oil cells which are common in Lauraceae. Specimen: SM.B Me 8375.

Family Leguminosae Juss. Legumes are represented at Messel by at least four different types of pods, ranging from small and single-seeded to long and multiseeded. Most are represented by only a small number of specimens and their affinity to modern legume groups is difficult to evaluate. It is likely that these represent fruits produced by the same plants as leaves and leaflets also preserved at Messel, among which five species have been recognised (Wilde 1989).

Genus Mimosites Bowerbank Mimosites spiegeli Engelhardt (Pl. 23a–f) 1922 Mimosites spiegeli Engelhardt, Abh. hess. geol. Landesanst. Darmstadt, 7 (4): 118, pl. 39, fig. 1.

Emended description: Pod stipitate (stipe up to 3.8 mm), one small calyx lobe observed (SM.B Me 19035, pl. 23c), peduncle up to 18 mm long. Pod elongate, usually curved (e.g. SM.B Me 7056, pl. 23b; 19035, pl. 23c), relatively thin (not woody), 6.0–11.2 cm long, 1.4–1.8 cm wide. Lateral 34

Comments: This species is readily distinguished from the other types of pods from Messel by its longitudinal curve, seeds oriented perpendicular to long axis of pod and by prominent transverse partitions delimiting the seed chambers. Specimen SM.B Me 19352 has less prominent chamber partitions than other specimens. We reillustrate here the type specimen of Mimosites spiegeli Engelhardt (pl. 23a). The species is now represented by many specimens in the Messel collections. As stated in the original description by Engelhardt (1922), the species is similar in pod size, morphology, and chamber delimitation to Mimosites browniana Bowerbank from the London Clay locality of Assington, Suffolk, England. Reid & Chandler (1933) reexamined the holotype of M. browniana, and noted that it was no longer possible to see finer details of structure and that therefore, they could offer “no opinion as to the generic relationship of the fruit” although Bowerbank’s original diagnosis stated “fruits which belong to the natural order Mimoseae.” Bowerbank’s specimen from the London Clay is refigured here (pl. 24h) for comparison. Although M. spiegeli could indeed be a mimosoid legume, there are few diagnostic characters that help to narrow down its affinities. Several species of Albizia, for example, have fruits that are similar in gross morphology (P. Herendeen, pers. comm., 2011). Specimens: Holotype HLMD-Me-1760. Other specimens SM.B Me 4014, 4671, 4884, ?4885, 7036, 7056, 7057, 7067, 7072, 7162, 19035, 19098, 19352, 19353, 20054.

Genus Leguminocarpon Goeppert Leguminocarpon herendeenii sp. n. (Pl. 24a–d) Diagnosis: Fruit stipitate, 1–2-seeded specimens, 14 mm long and 13.5 mm wide; multi-seeded specimen 19.5 mm long and 10.5 mm wide (excluding stipe); pedicel 3.7 mm, stipe up to 4.2 mm long. Calyx persistent with at least three lobes visible, Fruits relatively thick, asymmetrical, subcircular in single-seeded to oblong in multiple-seeded specimens, bounded by veins on both sides. Margins thickened around the periphery, one with a


groove. Apical protrusion different in prominence (mucronate in single seeded specimen to broadly conical in multiple-seeded specimen). Surface of fruit transversely rugulate to irregularly verrucate. Venation not visible. Seed size, shape, and outline unknown. Valve displacement suggests dehiscent. Etymology: This species name recognises the important contributions to our knowledge of fossil legumes made by Pat Herendeen. Comments: Few-seeded pods are not uncommon among legumes. Examples from several extant genera were illustrated by Herendeen & Crane (1992). Of those, Copaifera cordifolia Hayne (their fig. 16) looks most similar to our single-seeded specimens (e.g. SM.B Me 17519, pl. 24a), but that fruit differs in having oblique rugulae. Although we have designated a 2-seeded pod as holotype (pl. 24c), we have included single-seeded pods and a probable three-seeded pod (SM B Me 15470) because their texture appears similar and they preserve similar remnants of the perianth. One of the specimens shows one large and one aborted seed (pl. 24d). Herendeen (pers. comm. 2011) noted that these specimens resemble the pods of a caesalpinioid legume, and that some of the Asian species of Crudia have similarly small fruits, transverse oriented reticulations on the valves, and sometimes constrictions between the seeds. Specimens: Holotype SM.B Me 20422 (here designated, pl. 24c, showing two equally developed seeds, 3 calyx lobes visible). Paratypes SM.B Me 2303, 5635, 15453, 16859 (two-seeded), 17519, 18271 (one large apical seed and one small abortive basal seed), 21270 (one large seed and a small abortive seed at apex).

Leguminocarpon sp. 1 (Pl. 24e, f) Description: Pod, almost straight but with a slight curvature, relatively thin (not woody), length 125 mm, maximum width 18 mm, gradually narrowing from broadest point toward apex and ultimately narrowing abruptly toward an acute apex. Placental margin somewhat concave; opposite margin correspondingly convex. Placental margin with a thick band (almost 2 mm thick) from which the individual seed outlines arise. Opposite margin not thickened. Faint ridges separating seed chambers at right angles to the placental margin, extending only half way to the opposite side. Seed outlines pronounced, shape of seed outlines rounded-rectangular (seed width parallel to length of pod 5–6.5 mm, seed length 6–8 mm). Seed outline extending about half way to the non-placental side. Venation not visible. Comments: Gleditsia wesseli Web., as figured with a line drawing from Messel by Engelhardt (1922), shows a similar seed size and position but differs in other characters, such as what seems from the illustration to be

more strongly developed transverse partitions between seeds. Because the figured specimen could not be relocated, a more detailed comparison is impossible, and we cannot be sure that Engelhardt’s specimen is not another representative of his “Mimosites” spiegeli. A single specimen, SM.B Me 7057, has a thick placental vein, obvious placentas and short seeds (like this taxon) but also exhibits obvious chamber partitions. All other specimens fall clearly into one or other of the taxa as characterized above. Herendeen (pers. comm. 2011) considers it likely to be a caesalpinioid legume (but not likely to be Gleditsia), and noted that the seeds are larger than would be expected for Mimosoideae. Some extant species of Senna and of Bauhinia have fruits similar to this fossil. Specimens: SM.B Me 2323, 8693, 7035, 7055, 19800.

Leguminocarpon sp. 2 (Pl. 24g) Description: Fruit an elongate, straight pod, relatively thin, 65 mm long and 17 mm wide, parallel-sided, bounded by longitudinally striate narrow “wings” on both sides with wider band at the placental suture, 2 mm, and the narrow band on the dorsal margin, 0.8 mm. Apex and base rounded or obtusely pointed. Seed chambers narrow, 2–3 mm wide, with very fine partitions. Venation seen near apex of specimen, a reticulum of evenly sized fine veins. Seed outlines narrow elliptical, c. 3 mm wide, 7 mm long, oriented approximately at right angles to fruit axis and borne on long funicles. Comments: This species is known from a single specimen broken at the base. It is distinguished from the other Messel legumes by the very narrow seed chambers. The attenuated margins of the pod indicate that it might represent an indehiscent pod. Most Gleditsia species have similar pod margins, but not the narrow seed chambers seen in this fossil. Specimen: SM.B Me 23628.

Family Lythraceae J. St.-Hil. Genus cf. Decodon J. F. Gmel. (Pls 25a–g, 26a–g) Description: Paniculate infructescences with fruits borne on elongate peduncles c. 6 mm long. Peduncle articulated about 1–2 mm below the fruit with remnants of bracts or other abortive flowers. Individual fruits globose, 3.5–6 mm in diameter, with a single style persisting at the apex (pl. 25d). Fruits composed of closely packed seeds (4–6 visible) that are triangular to polygonal in exposed outline. Seeds 1.5–2.5 mm in diameter, wedge shaped with narrow ends directed toward the centre of the fruiting head, facetted laterally through mutual compaction. 35


Individual seeds 1–1.2 mm diameter near surface of the fruit, convex rounded distally, tapering proximally. SRXTM digital sections reveal elongate, asymmetrically curved embryo cavity, occupying small proportion of the seed volume. Main volume of seed composed of isodiametric cells with thin walls but both the outer wall of seed coat, and a wall delimiting the embryo chamber, are composed of smaller thick-walled cells forming an X-ray dense tissue (SM.B Me 20157, pl. 26b–g).

Comments: A single fruiting axis shows the characteristic organization of Magnolia. Although Magnolia has been identified from the Early and Middle Eocene of Europe previously on the basis of seeds, this specimen shows that the morphology of the fruit was also consistent with that of the extant genus.

Comments: One of the specimens clearly shows the infructescence and branching pattern (pl. 25a, b) and another is a single fruit on a peduncle with persisting single style (pl. 25d). Isolated clusters of seeds are also found, apparently representing the fruit with its thin pericarp deteriorated (e.g. pl. 25e, f). Some fruits are exposed in surface view with the seeds apparent as bulges (pl. 25b, d), others are fractured longitudinally and/or obliquely to reveal the seeds in lateral view (pl. 25c). The seeds, visible both in surface view and in lateral view in broken fruits, are wedge shaped by mutual compaction within the globose fruit, but are rounded on the distal surface. In addition to viewing fruits exposed in random fractures, we examined a complete seed cluster by SRXTM (pl. 26a–g) to reveal internal organization and seed anatomy. Virtual sections of the specimen in multiple directions indicate that the seeds contain a longitudinally curved embryo chamber plus a large area composed of parenchymatous tissue (pl. 26f, g). The specimens are very similar to fruits and seeds of modern Decodon (pl. 26h–j) which is endemic to eastern North America today. The anatomy of the Messel fossils as revealed by SRXTM compares closely with that of fossil Decodon described by Cevallos-Ferriz & Stockey (1988) using permineralised material from the Middle Eocene Princeton Chert. In addition to the Princeton chert occurence, Decodon-like infructesences and fruits are known from the Eocene of Oregon, as well as the Miocene of the Czech Republic (Kvaček & Sakala 1999), with seeds described from many localities in Europe, North America and Japan (Matsumoto et al. 1997).

Magnolia spp. (Pl. 27a–f)

Specimen: HLMD-Me-15680.

Description: Seeds obovate to transverse-elliptic with smooth surface, length and width range from 6–8 mm (l/w 0.75–1.3). Seeds sometimes pointed toward micropylar area, chalazal surface more or less truncate to slightly rounded. Chalaza shallow, more or less circular (0.2–0.3 mm in diameter). Seed coat formed of anticlinally oriented columnar cells. Comments: Collinson (1988: tab. 1) referred to two genera of Magnoliaceae from Messel based on ten specimens. On re-examination only the specimens listed below show diagnostic characters of the family. Following Collinson (1983a), who noted that an adequate analysis of intraspecific variation among fossil Magnolia seeds is needed before distinguishing the taxa from the London Clay flora, we have refrained from assigning formal epithets to the Messel material. Specimens: SM.B Me 4757, 7751, 10510, 10611, 20156, 21383, 21510, 21515. The listed specimens are examples with the chalaza facing upward from the shale. Specimens with the chalaza oriented downward in the shale cannot be as confidently identified, because they may resemble unrelated disseminules of similar shape.

Family Mastixiaceae Calest. Mastixiaceae are one of the more diverse families at Messel. At least four taxa are present and one, Eomastixia, is one of the more abundant fossils in the collections.

Specimens: SM.B Me 2056, 2095, 2103, 7539, 7075, 17917, 20157.

Family Magnoliaceae Juss. Genus Magnolia L. Magnolia sp. 1 (Pl. 27g) Description: Ovoid multifollicular fruit 15 mm high and 13 mm wide, with stout peduncle that is 5 mm wide near the base, and widens to 9 mm at scars of perianth and stamens below the follicles. Follicles closed, spirally arranged. Approximately 15 follicles visible on exposed surface, estimate 30 in total, follicles subelliptical, 5 mm wide, 9–12 mm in long, 2–2.5 mm thick. 36

Genus Eomastixia Chandler Eomastixia cf. rugosa Chandler emend. Chandler (Pls 28a–m, 29a–g) Description: Fruit, syncarpous, ovoid to subglobose, 9–18 mm high, 8–15 mm wide, rounded basally and rounded to pointed apically, with an apical disk (SM. B Me 2241, van Aarssen et al. 1994) inferred to mark the apical area lacking perianth. Surface longitudinally sinuously striate, at least 5–6 striae on exposed surface (indicating at least 10–12 on complete fruit). Surface with irregular topography with tendency to longitudinal rugulae. When abraded, densely spaced, longitudinally

elongate resin bodies are exposed. In transverse fracture (n = 3), and digital section the outermost fruit wall has a slightly different texture to the inner and contains many large and often juxtaposed resin pockets (pl. 28c–e). Inner portion of fruit composed of radiating fibres swirling around the locules. Two, four or five U- or V-shaped locules per fruit, each with a dorsal infold and sometimes a large resin pocket up to 1.7 mm diameter situated in fruit wall on same radius as the infold. No axial (central) vascular bundle present, other vasculature not evident. In longitudinal fracture (n = 2; van Aarssen et al. 1994: fig 2b and digital sections of SM.B Me 2270) resin occurs in discrete longitudinal bands close to the surface up to 0.6 mm in diameter. Comments: Specimens are usually compacted laterally (pl. 28a–i), such that transverse sections show very compressed, distorted locules (pl. 28c–d), but occasionally specimens are compacted more or less vertically, so that transverse fractures and sections show the number and arrangement of locules more clearly (pl. 28k–m, pl. 29a–g). The number of locules ranges from 2 to 5 (2: pl. 29d–f; 3: pl. 29c, g; 4; 5: pl. 29a, b). Externally the fruits do not reveal the sulci of infolds, and there is no obvious expression of germination valves. This species is readily recognisable among the ellipsoidal fruits from Messel by the longitudinal striations and commonly exposed subsurficial yellowish resin masses. The fruits were treated as undetermined large ovoid fruit with resin ducts by Collinson (1988) and provisionally referred to as Eomastixia aff. rugosa by van Aarssen et al. (1994). This species corresponds closely to the genus Eomastixia Chandler 1925/26 emended 1962 in the bi- to multilocular fibrous endocarps with U or V-shaped locules, and prominent resin cavities and/or ducts towards the periphery of the fruit (Chandler 1925/26, 1961a, 1962; Mai 1993). Germination valves are poorly defined, not obvious, as in the type material from Hordle. The Messel material is distinguished from the Hordle species by having a much larger number of resin bodies preserved, especially close to the surface (pl. 28c–e). Although occasionally (e.g. SM.B Me 8895 when cut) resin may be absent. Two- to three-loculed specimens are typical at Hordle, but Hóly (1975) provided an addendum to the generic diagnosis, indicating a range of 1–4 locules based on two additional species from the Bohemian Tertiary. The Messel population ranges up to 5 locules (pl. 29a, b). Bilocular specimens are less common, and the example we illustrate here (pl. 29d, e) has much less resin than the multilocular specimens. The resin chemistry was studied by van Aarsen et al. (1994) and shown to be a sesquiterpenoid type previously thought to be characteristic of Dipterocarpaceae. This indicates that sesquiterpenoid compounds have a much wider botanical source than was previously recognised. The only other fossil mastixioid known with large quantities of resin is Retinomastixia Kirchh. from the Miocene.


That genus has only unilocular fruits, and has the resin pockets throughout, rather than confined to the external portion of the fruit. Specimens: Transversely fractured SM.B Me 4053 (5-loculed, transverse), 5174, 20153 (4-loculed, transverse); 11280 (3-loculed), 15448 (2-loculed); longitudinally fractured 5172, 5175. Other specimens 2229–2245, 2247–2249, 2250–2260, 2266–2271, 2319, 2576, 2664– 2666, 2672, 4015, 4039, 4048, 4053, 4054, 4059, 4070, 4100, 4127, 4140, 4154, 4160, 4235, 4395, 4396, 4398, 4400–4402, 4653, 4701, 4797, 4800, 4801, 4804, 4887–4897, 5083, 5084, 5173–5192, 7154, 7593, 7594, 7602, 7172, 7173, 7240, 7220–7222, 7206, 7208, 8267, 8895, 18072, 19071, 20150.

Genus Mastixia Blume ?Mastixia sp. (Pl. 27h–t) Description: Endocarp oblong-ovoid, rounded to truncate basally and apically, smooth, with a prominent dorsal furrow (1–2 mm wide) which extends almost the full length of the endocarp. Endocarp 8–11 mm long, 3.5–5 mm wide. Centre of dorsal furrow occupied by a longitudinal ridge (e.g. SM.B Me 614, pl. 27j; 8359, pl. 27h). Ventral side rounded. Germination valve not observed. Comments: We considered the possibility that these specimens are seeds of phoenicoid palms. In phoenicoid palms (e.g. Phoenix L.) the longitudinal groove flares at one end where it does not reach the tip of the endocarp and indents the endocarp as it passes over the opposite end. Specimen SM.B Me 19992 (pl. 27k) is the only Messel specimen with this appearance, although the indented end may be damaged and it lacks the longitudinal ridge (which might be a raphe ridge if this was a phoenicoid palm seed). SRXTM of this specimen yielded little additional information (pl. 27l–s) and failed to reveal any information concerning the possibility of a dorsal scar (such as demonstrated for Miocene Phoenix seeds by Bůžek (1977). The specimens are similar to Mastixia parva Reid & Chandler in the morphological characters available (Reid & Chandler 1933, Collinson 1983a: fig. 127), but some specimens of M. parva have rugose endocarps (Chandler 1961b). They are also similar in size, morphology and smooth surface to specimens from the Clarno Nut Beds treated as Mastixia sp. (Manchester 1994). Direct comparison is hindered, however, because the Messel material is compressed, whereas the London Clay and Clarno specimens are permineralized. Overall, therefore, we favour the interpretation of these specimens as Mastixia – but recognise that we cannot rule out other affinities. Additional new specimens may provide options for preparation to release them from the oil shale for further investigation. Two Messel specimens show a surrounding halo of softer organic tissue (SM.B Me 23368, pl. 27l; SM.B Me 20574, pl. 27i) that could be remains of a fleshy outer layer, if not a coprolite. 37

Collinson, M., Manchester, S. & Wilde, V.: Fossil Fruits and Seeds of the Middle Eocene Messel biota, Germany Specimens: SM.B Me 306, 614, 2174–2176, 8359, 19992, 20574, 23368.

Genus cf. Beckettia Reid & Chandler (Pl. 29h–l) Description: Fruit/endocarp ovoid, 12–14 mm long, 9.3–10 mm wide, with a prominent elongate germination valve visible on one face; possibly another inferred to be on the opposite side. Valve margins narrow to pointed apex and broaden to wide base. Exposed surface of valve with a median, longitudinally striate ridge bordered by smooth marginal bands that extend from base. Surface of the fruit outside germination valve smooth to wrinkly. Internal structure not observed. Comments: The elongate germination valves visible on the surface of these specimens support the placement in Mastixiaceae and they are particularly similar in configuration to those of Beckettia Reid & Chandler 1933 from the London Clay (e.g. pl. 29m). Like Beckettia, these fruits do not possess resin cavities. The flattening of the Messel specimens suggests a non-rigid composition, as would be expected for compressed specimens of Beckettia which is known from pyritised specimens to be composed largely of parenchymatous tissue between the locules. Specimens: SM.B Me 2663, 4118, 11183, 17531, 23139.

Indeterminate mastixioid (Pls 29n, o, 30j, k) Description of SM.B Me 2264: Fruit oblong ellipsoidal, irregularly longitudinally ridged. Two prominent longitudinal grooves on opposite sides of the endocarp mark infolds in the germination valves. Valves elongate oval, running the full length of the endocarp, narrowing evenly to both ends, and wide (5.9 mm), almost the full width of the fruit. Whole endocarp 6.8 mm wide. Thickness in plane of compression 3 mm, indicating substantial original thickness of sclerified tissue. A thin outer cuticle preserved in patches; the outermost part of the endocarp wall includes patches of periclinally, longitudinally elongate cells; the main thickness composed of small isodiametric cells up to 90 μm in diameter). Internal organization unknown, inferred to be bilocular on basis of two opposite germination valves. Comments: The very wide and evenly narrowing germination valves distinguish SM.B Me 2264 from other Messel mastixioids. We refrain from naming this taxon as the internal structure is not known and the conspecificity of the three specimens is not proven. SM.B Me 2225 and 2226 are approximately twice the size of SM.B Me 2264 (both being c. 12.5 mm wide), and their germination 38

valves are much less clearly defined. They may or may not belong to the same taxon. Specimens: SM.B Me 2225, 2226, 2264.

Genus Mastixiopsis Kirchheimer Mastixiopsis nyssoides Kirchheimer (Pl. 30a–i) 1936 Mastixiopsis nyssoides Kirchheimer, Beih. Bot. Centralbl., 55: 291–292, pl.7, fig.5a–g.

Description: Fruit ellipsoidal, 17–22 mm long, 11– 13 mm wide, with c. 10 straight longitudinal bundles on the surface. Thick walled, unilocular. Fruit wall composed of fibers and/or sclereids that are anticlinally oriented in the outer 1/4 of the wall, and more isodiametric sclereids in central portion. Locule U-shaped in cross section, with transversely striate lining. Germination valve elongate and wide. Resin apparently absent. Comment: This species is distinguished from the other Messel mastixioids by its thick woody wall and lack of resin. We made first-hand comparisons with the type material of Mastixiopsis nyssoides from the lower Eocene of Riestedt, Central Germany, and with the specimen from England attributed to this species by Mai (1993) from the Dorset Pipe Clays at Arne that Chandler (1962) initially attributed to Mastixia cantiensis Reid & Chandler. The Messel specimens also show widely spaced straight longitudinal bundles like the type material, and the thickwalled, unilocular construction. The infold contains two grooves. No resin was observed in these fossils; in the Riestedt material, only scattered rare small resin cavities are seen (Manchester, pers. obs.). Specimens: SM.B Me 5082, 5739, 7176, 23283.

Family Menispermaceae Juss. Menispermaceae are the most diverse family recognised in the Messel fruit and seed assemblage, including at least 14 genera. Traditionally, eight tribes are recognised in the Menispermaceae on the basis of fruit morphological characters (Diels 1910), but recent molecular work indicates that some of the genera must be regrouped, and that some modification to the traditional tribe scheme is necessary for the classification to circumscribe truly monophyletic groups (Wang et al. 2007, Hoot et al. 2009, Jacques et al. 2011). In this treatment, we make reference to the traditional tribal classification because it applies well to fruits, the organs that we are considering here, but reference is also made to the more recent works because the classification is still in flux as increased sampling for both molecular and morphological analyses are needed. Morphologically, the Messel menisperms fall


mainly into two basic groups, those with straight, boat shaped endocarps, and those with bent, C- or U-shape. The boat shaped fruits mostly represent the Tinosporeae and/or Fibraureae tribes sensu Diels. Ortiz et al. (2007) recognise an “expanded Tinosporeae” including most of Diels's Fibraureae and all his Peniantheae, as well as genera traditionally treated as Tinosporeae. Although these authors excluded Tinomiscium Miers results from other molecular studies (Hoot et al. 2009, Jacques et al. 2011) indicate that it belongs within the expanded Tinosporeae.

Genus Diploclisia Miers Diploclisia rugulosa sp. n. (Pl. 31a–d) Diagnosis: Endocarp bisymmetric, dorsiventral length 10.5–12.5 mm, width 8.2–8.5 mm (n = 3), laterally flattened, obliquely obovate in lateral view with broad rounded dorsal surface, with a dorsal crest (“keel”) running around the plane of bisymmetry, base acutely rounded. Smooth central condylar depression asymmetrically elliptic and slightly raised. Central area surrounded by a slightly asymmetric horseshoe-shaped raised ornamented area (lateral crest). One limb of the endocarp slightly longer than the other. Central area lacks obvious foramen, but contains elongate slit in a more or less median position, extending two thirds the length of the endocarp from the base. Lateral crest area ornamented by about 30 conspicuous radiating irregular nodular ridges which sometimes extend across the dorsal crest, but do not protrude out of the profile of the endocarp and do not bear spines. Etymology: This species name refers to the ornamentation of the fruit. Comments: Genera with horseshoe-shaped endocarps were formerly treated in tribe Menispermeae (syn. Cocculeae), but recent molecular studies indicate that they occur in five different clades: Diploclisia, the SinomeniumMenispermum clade, the Perycampylus-Sarcopetalum clade, Cocculus, and the Stephania-Cyclea-Cissampelosclade. All of these have C- or U-shaped endocarps, with a smooth central, condylar area and a regular pattern of ornamentation over the dorsal and/or lateral crests. Diploclisia, which has two extant species in Asia, is distinguished by elongate “hairpin” configuration (pl. 31n, Jacques 2009), with about 30 transversely oriented short ribs expressed on the distal margins of the lateral crests that do not reach over the dorsal crest. A central slit extends from the base toward the central part of the curved endocarp, but the absence of a foramen through the condyle distinguishes this genus from several other extant genera having horseshoe-shaped endocarps. In comparisons of fossils with C- and U-shaped endocarps

with extant taxa, we have found the comparative table compiled by Liu and Jacques (2010) to be very useful. The two extant species differ significantly in size e.g. 4.4 mm long in D. affinis Diels, vs. 19 mm long in D. glaucesens Diels (Jacques 2009). The Messel species differs from both the English and North American Eocene species. D. bognorensis Chandler 1961b (London Clay) is smaller (6.69 mm long, 5.5–6.5 mm wide) and has a distinct central foramen (unlike the extant species) and less nodular ridges, but with a hint of poorly preserved spines protruding from the periphery of the endocarp. The North American species, D. auriformis (Hollick) Manchester (1994) is also smaller (length 6.3–8.0 mm, width 4.0–6.8 mm), with radiating ribs that are not nodular, and it possesses a central foramen. Specimens: Holotype SM.B Me 2069 (here designated, pl. 31b). Paratypes SM.B Me 4090, 4912 (removed from oil shale), 5155, 10481, 17539, 17713, 17772, 20539.

Genus Stephania Loureiro Stephania hootae sp. n. (Pl. 31e–h) Diagnosis: Endocarp bisymmetric, dorsiventral length 7.0–7.6 mm, width 5.0–6.4 mm (n = 4), laterally flattened, obliquely obovate in lateral view with broad rounded dorsal surface, with a dorsal crest (“keel”) running around the plane of bisymmetry; base obliquely truncate. Central condylar depression obpyriform, smooth and more or less flat. Central area with distinctive oval to circular foramen central to the endocarp, and a slit extending from foramen to base. Central area surrounded by a slightly asymmetric horseshoe-shaped raised ornamented area (lateral crest). One limb of the endocarp distinctly longer and protruding at the base when fully preserved. Lateral crest surmounted by c. 30 conspicuous spines. A corresponding row of spines located along dorsal crest at periphery of the endocarp; these spines sometimes connected to spines on the lateral crest by shallow ridges. Etymology: The epithet, hootae, recognises the contributions of Sara B. Hoot in understanding relationships among extant Menispermaceae. Comments: These endocarps differ from Diploclisia rugulosa sp. n., above, by the smaller size, prominent spines on both lateral and dorsal crests, more oblique base and in the presence of a central foramen. We are not aware of spiny ornamentation in extant Diploclisia although the general form of the endocarp is similar. Very similar endocarps with the same pattern of spiny ornamentation, on both lateral and dorsal crests, a smooth condylar area and an elliptical central foramen occur among extant species of Stephania, particularly 39


S. hernandifolia (Willd.) Walp. (pl. 31o), and S. glabra Roxb. (Miers) as illustrated in Jacques (2009: fig. 11c). S. palaeosudamericana, recently described from the Paleocene of Colombia (Herrera et al. 2011) differs by the absence of conspicuous dorsal spines. Specimens: Holotype SM.B Me 2070 (here designated, pl. 31f). Paratypes SM.B Me 2071, 2072, 2660, 4645, 4680, 5154, 5156, 5161, 5749, 7583, 8307, 8665, 12253 (now broken), 20538, 20540, 20541, 24972.

Indeterminate Menispermeae (Pl. 31i) Description: Endocarp bisymmetric, dorsiventral length 4.3 mm, width 3.0 mm (n = 1), laterally flattened, obliquely obovate in lateral view with broad rounded dorsal surface, with a dorsal crest (“keel”) running around the plane of bisymmetry; base obliquely truncate, with a prominent slit-like sinus extending to the centre of the endocarp. Margin of sinus raised. Central area surrounded by a slightly asymmetric horseshoe-shaped, lateral crest, ornamented with a row of c. 30 rounded mounds. One limb of the endocarp distinctly longer and protruding at the base. Comments: This specimen is similar in gross form to Diploclisia sp. described above, but is less than half the size. It may represent another species of the same genus, but would be smaller than either of the two living species. Specimen: SM.B Me 16573.

Genus Martinmuellera gen. n. Martinmuellera tuberculata gen. et. sp. n. (Pl. 31j–m) Diagnosis: Endocarp bisymmetric, dorsiventral length 5–7 mm, width 5.7–6 mm (n = 2), laterally flattened, more or less circular in lateral view but slightly asymmetrical, with broad rounded dorsal surface, dorsal crest (“keel”) absent. Base of endocarp obliquely truncate. Central condylar depression on each lateral surface more or less subcircular, smooth, and nearly flat. No obvious foramen in the central area but a slit extends from the base to the centre and slightly displaced to the longer limb of the endocarp. Central area surrounded by a very narrow C-shaped lateral crest. Lateral margins beyond the lateral crest densely ornamented by discrete, closely spaced, irregular to conical tubercles, sometimes irregularly arranged, or arranged in rows. There are about five tubercles across the radius and 26–28 around the circumference. Tubercles are 0.2–0.5 mm broad. Etymology: This generic name recognises M artin Müller, for his many years devoted to the laboratory preparation of Messel specimens including those utilized 40

in this investigation. The specific epithet refers to the surface ornamentation. Comments: The C-shape of the endocarp is consistent with the Menispermaceae, but we are unaware of extant or other fossil species with this kind of sculpture. Although tubercular ornamentation occurs in some extant species such as Pericampylus glaucus (Lam.) Merr. (pl. 31q), Cocculus carolinus Hort. (pl. 31r) and Cissampelos pareira L. (Forman 1986: fig. 17), the arrangement and shape of the tubercles differs. Some features of the locule can be observed through fractures in SM.B Me 4916 (pl. 31j). The cluster of endocarps shown in pl. 31j shows no regular organization and is, therefore, likely to be derived from a coprolite. In contrast the cluster in pl. 31k shows limbs of the endocarps pointing inwards in most cases, and a more regular arrangement of the endocarps, suggestive of the organization of fruits in modern Menispermaceae fruiting branches. Specimens: Holotype: SM.B Me 8555 (here designated, pl. 31l). Paratypes SM.B Me 229, 4439, 4768, 4916 (cluster), 7204, 7246, 12305 (cluster) 16442, 17404 (cluster), 23266. SM.B Me 20545 might belong to this taxon but has a raised and radially ridged concentric ridge around the central area.

Tribe Menispermeae DC. Genus Palaeosinomenium Chandler The genus Palaeosinomenium was established by Chandler (1961b) to accommodate fossil endocarps “closely comparable with Sinomenium Diels and Menispermum L., but with more oblique endocarp and elongate foramen”. These might be regarded as relatively minor characters for generic distinction and Mai (1997) considered that the characters used to diagnose Palaeosinomenium (and Wardensheppeya Eyde) are really only suitable at the species level. Nevertheless, we retain Palaeosinomenium as a useful morphogenus for endocarps whose affinities might lie with either Sinomenium or Menispermum or both. Sinomenium and Menispermum are sister taxa in the molecular phylogeny of Hoot et al. (2009) and are very similar in endocarp morphology both being asymmetrically C-shaped with a small elliptical foramen, one lateral crest on each side with fine radial ribs except over the condylar region. However Liu & Jacques (2010) summarized endocarp characters that can distinguish these two genera, including a more circular endocarp outline and a more prominent, angular ventral notch in Menispermum, higher number of radial ridges on the dorsal and lateral crests in Menispermum (27–39) than in Sinomenium (18–21). Kirchheimer (1957) also noted that the radial ribs are weaker in Menispermum than those in Sinomenium. These Messel fossils do not precisely fit one of these genera to the exclusion of the other, so we place them in the morphogenus Palaeosinomenium. If further work on the extant representatives of this clade leads to the sinking of Sinomenium within Menispermum, as the molecular data


may suggest, then the older name Menispermum would take priority and these fossils would fit well within a broadened concept of the extant genus.

Palaeosinomenium ornamentum sp. n. (Pl. 32b–d) Diagnosis: Endocarp bisymmetric, dorsiventral length 4.5–6.0 mm, width 5.7–6.6 mm (n = 3), laterally flattened, reniform in lateral view with broad rounded dorsal surface, with a dorsal crest (“keel”; equals marginal flange sensu Chandler 1961b) running around the plane of bisymmetry, base concave to convex but mostly almost straight. Smooth central condylar depression almost subcircular to obovate, asymmetrical and more or less flat. Central area has a distinctive elongate foramen close to the basal (ventral) side (foramen about a quarter of the dorsiventral length of the endocarp). Central area sometimes raised around the foramen. Central area surrounded by a slightly asymmetric horseshoe- to Cshaped lateral crest, on the top of which there is always a narrow groove. To the inside of the lateral crest there are inconspicuous nodules in a single row in some specimens, apparently absent in others. To the outside of the lateral crest is a row of about 15–20 radially elongate ridges (sometimes club-shaped, sometimes branching) extending slightly more than half the distance between the crest and the periphery of the endocarp. Between these ribs and the periphery the endocarp is ornamented with closely spaced radially elongated finer ribs giving it a striated appearance (rarely they may be absent, rarely more strongly developed as ridges opposite ribs. Etymology: The specific epithet refers to the obvious ornamentation. Comments: This species closely resembles that known from Eckfeld (Neuffer et al. 1996: fig. 68). Collinson (1996: 193–194) summarized characteristics of British Paleogene Palaeosinomenium. P. venablesii Chandler 1961b from the Bracklesham Group (and Messel, see below) is similar to this species but lacks striations on the outside of the endocarp. Furthermore, the coarse ridges are not club-shaped in P. venablesii and the Bracklesham specimens are also smaller. Indeed all British Palaeosinomenium are smaller than this species. Specimens: Holotype SM.B Me 7343 (here designated, pl. 32b); Paratypes SM.B Me 2074, 2075, 2077, 4913, 7212, 8425, 17895, 18001, 20547, 20548.

Palaeosinomenium venablesii Chandler (Pl. 32e) 1961 Palaeosinomenium venablesii Chandler, Lower Tertiary Floras of Southern England, 159–161, pl. 16, figs 9–13.

Description: Endocarp bisymmetric, dorsiventral length 3 mm, width 3.5 mm (n = 1), obliquely C-shaped with a broadly rounded dorsal margin and truncate ventral margin, with a dorsal crest (“keel”) running around the plane of bisymmetry, base almost straight, slightly damaged. Central condylar depression subcircular, smooth, and almost flat with an oval foramen near the longer endocarp limb. Central area surrounded by a C-shaped lateral crest to the outside of which is a row of about 18 radially aligned ridges extending most of the distance to the endocarp periphery, in some cases continuing to the edge of the dorsal crest. Comments: This single specimen differs from Palaeo sinomenium ornamentum, described above, in much smaller size and less pronounced ornamentation, lateral ridges not expanding into a club-shaped morphology. It also differs from P. sp. described below in smaller size and less pronounced ornamentation on the endocarp periphery. It corresponds closely to the type species of Palaeosinomenium from the early Eocene of England (Chandler 1961b, 1978; Collinson 1996), which has also been recognised from the Middle Eocene of Oregon (Manchester 1994) and Huadian (Manchester et al. 2005). Specimen: SM.B Me 14061.

Palaeosinomenium sp. (Pl. 32f–i) Description: Endocarp bisymmetric, dorsiventral length 5.25–6.25 mm, width 4.75–6.25 mm (n = 4), laterally flattened, reniform in lateral view with broad rounded dorsal surface, with a dorsal crest (“keel”) running around the plane of bisymmetry; base strongly asymmetrical, concave with one distinctly longer limb (longer limb distinctly pointed). Central condylar depression on each lateral face, semicircular, smooth, flat, sometimes with slightly raised areas. Central area has indistinct elongate foramen close to the basal (ventral) side (foramen less than a quarter of the dorsiventral length of the endocarp). Central area surrounded by an asymmetric C-shaped, lateral crest, on the top of which there is a discontinuous inconspicuous narrow groove. Lateral crest ornamented with up to 25 relatively narrow radially elongated short ridges which continue across the dorsal crest and sometimes protrude from the periphery of the endocarp. To the inside of the lateral crest on some specimens (SM.B Me 4011, pl. 32f; 4193, pl. 32g) there are conspicuous inwardly radiating nodules but these are lacking on SM.B Me 2076. Comments: This species differs from P. venablesi in having larger size and more pronounced ornamentation, especially on the periphery of the endocarp and dorsal crest. P. ornamentum sp. n. has pronounced club shaped or branching ridges adjacent to the lateral crest, which 41


contrasts with P. sp. where the ornamentation is very similar across the endocarp consisting of narrow elongate radial ridges. Specimens: SM.B Me 2073, 2076, 4011, 4193, 17855.

Genus Pericampylus Miers ?Pericampylus sp. (Pl. 32j) Description: Endocarp bisymmetric, dorsiventral length 2.5 mm and 3 mm wide (dimensions including spines), laterally flattened, reniform in lateral view with broad rounded dorsal surface, with a dorsal crest (“keel”) running around the plane of bisymmetry; base oblique with one longer limb. Central condylar depression obovate, asymmetrical, smooth and more or less flat with a short slit near the longer limb but no obvious foramen. Central area surrounded by a horseshoe shaped lateral crest on the top of which is a narrow groove. To the inside of the lateral crest there are very minor irregular nodules in a single row. To the outside of the lateral crest there are 19 radially elongated broad ridges extending nearly to the periphery of the endocarp. Opposite each of these on the dorsal crest is a stout spine, extending radially beyond the endocarp profile, up to 0.5 mm long and up to 0.15 mm wide, tapering to a blunt tip. Comments: The very small size and dorsal crest with prominent spines distinguish this species from other Messel Menispermaceae. It is similar in form and ornamentation to extant Pericampylus glaucus (Jacques 2009: fig. 9J), but is smaller and more strongly asymmetrical. Specimen: SM.B Me 23272.

Genus Wardensheppeya Eyde Wardensheppeya sp. (Pl. 32a) Description: Endocarp bisymmetric, dorsiventral length 5.5 mm, width 5 mm, laterally flattened, reniform in lateral view with broad rounded dorsal surface, with a dorsal crest (“keel”; equals marginal flange sensu Chandler 1961b) running around the plane of bisymmetry, base slightly oblique and slightly concave. Central condylar depression obovate, with raised patches, almost symmetrical. Foramen not evident. Central area surrounded by an almost symmetrical horseshoe-shaped lateral crest on the top of which there is a narrow groove. To the inside of the lateral crest is a single row of irregular nodules. To the outside of the lateral crest is a row of 16 radially elongated nodular ridges extending nearly to the periphery of the endocarp. Endocarp smooth between these ribs and the periphery. 42

Comments: Wardensheppeya was distinguished (as Wardenia) by Chandler (1961b, 1978) and by Collinson (1983a) from Palaeosinomenium in having a more nearly symmetrical endocarp. The limbs of the “horseshoe” are more or less equal and the central area almost symmetrical (i.e. “less obliquely curved” sensu Chandler 1961b) in contrast with the asymmetrical condition in Palaeosinomenium. (See also comments on genus Palaeosinomenium). The single Messel specimen is similar to W. davisii (Chandler) Eyde (1970) from the London Clay (see also Chandler 1978), and from Le Quesnoy (Jacques & De Franceschi 2005), but is distinguished by the much more elliptic shape of the central area and the coarse ribbing extending most of the distance between the lateral crest and the periphery of the endocarp. Wardensheppeya poolensis Chandler 1962 is the same size as the Messel specimen and the radial ridges are similar in morphology and in their extent on the endocarp. However, the central area is much broader and more subcircular than in the Messel specimen. W. marginata Chandler 1963 is much smaller (maximum dimension 3.5 mm) and the radiating ridges are more variable in length. Specimen: SM.B Me 2078.

Genus Karinschmidtia gen. n. Karinschmidtia rotulae gen. et sp. n. (Pl. 33a–o) Diagnosis: Fruit circular to subcircular shape (lateral view), maximum diameter 23.4–29 mm. Preserved short straight area with a possible pedicel scar, from which longitudinal strands extend into endocarp sinus. Adjacent to the probable pedicel scar, the junction between the flat and rounded margin appears to represent the base of the style. Pericarp (outer cuticular envelope) commonly preserved, wrinkled and folded during compaction. Cuticle (proven by fluorescence) resistant, reflecting a polygonal pattern of epidermal cells without stomata. Cuticle seen on broken specimens to envelope both lateral faces of the endocarp. Beneath the pericarp cuticle are dense dark striae, possibly representing fibres, that usually radiate from the periphery of the central area of the endocarp, but we have no evidence for attachment. Striae over the central area. Endocarp bisymmetric, horseshoe-shaped, dorsiventral length 11.1–13 mm, width 10.5–13 mm, laterally flattened, subcircular to slightly obovate in lateral view with broad rounded dorsal surface, with a pair of concentric horseshoe-shaped lateral crests on both lateral faces, running around plane of bisymmetry. Lateral crests surrounded by a single dorsal crest from which a winglike structure extends. Base oblique with prominent sinus on isolated endocarps. Obpyriform smooth central condylar depression, more or less flat. Central area smooth with no obvious foramen. Dorsal crest and paired lateral

crests equidistant from each other. One limb is slightly longer than the other. The endocarp lacks nodules, spines or radial ornamentation apart from the wing-like fibrous flange on the dorsal surface. Etymology: The genus name recognises the important contributions of Karin Schmidt in the curation of palaeobotanical collections and invaluable help with the citation of Messel specimens cited in this publication. The epithet rotulae (L, pertaining to wheel) refers to the wheel-like shape of the nearly circular endocarp. Comments: These endocarps are quite unlike any others found at Messel or known in the palaeobotanical literature. The horseshoe-shaped endocarp body is positioned in the centre of an almost entirely encircling flattened disk with fibrous spokes radiating as in a wheel. This is the only menispermaceous fruit type at Messel that has tisssues external to the endocarp so consistently preserved, including the pericarp cuticle. We considered whether the distinctive circular halo surrounding the endocarps in these specimens might represent a wing that was in the plane of bisymmetry on the dorsal surface or simply the flattened remains of fibrous fleshy mesocarp tissue of the drupe. Wrinkling and folding in the cuticle (e.g. SM.B Me 517 and 2000) of the dorsal surface is suggestive of a compacted fleshy fruit rather than a solitary wing as is the presence of the pericarp cuticle on both faces of the endocarps (seen on broken specimens). However, the prominent radiating fibres are generally absent over the central area of the endocarp (though present in SM.B Me 2000) in the fruits and some specimens [e.g. SM.B Me 20452 (pl. 33d, e) and 8553 (pl. 33g)] show that the tough radiating strands have terminal branching at the circumference of the specimen supporting the wing interpretation. Additionally, specimens SM.B Me 4241 and 24310 (pl. 33n) show areas of pericarp cuticle external to the fused margin of the radiating fibre strands. We therefore interpret these specimens as having a wing-like extension of the endocarp, in the plane of bisymmetry that was surrounded by softer mesocarp tissues and a resistant epicarp cuticle. Among extant Menispermaceae, fruits of Legnephora Miers (pl. 33p–r; Forman 1972: fig 1; Jacques 2009: fig. 7I) exhibit similar morphology with a wing-like extension of the endocarp in the plane of bisymmetry. However, modern species of this genus, including those illustrated by Forman, show just a single prominent lateral crest on each side of the endocarp (pl. 33p, q) but Karinschmidtia has two or three concentric lateral crests each side of the endocarp (pl. 33a–j). Also, the lateral crests are variously fimbriate in the species descriptions of Forman, but Karinschmidtia shows only smooth crests. As the number of lateral crests and their ornamentation tends to be a consistent feature within the species of various extant menispermaceous genera (Jacques 2009), this fossil does not appear to belong within Legnephora.


More detailed comparative work is needed to determine whether Karinschmidtia and Legnephora represent the same intrafamilial clade, or may represent parallel or convergent evolution. Three isolated endocarps (e.g. pl. 33h), lacking the protruding radiating fibers and flanging wing, have been included in the above description. They correspond in morphology and dimensions to those observed within the fruit. Specimens: Holotype SM.B Me 21233 (here designated, pl. 33c, m). Paratypes SM.B Me 471, 517, 2000–2003, 2090, 4241, 4728 (with possible pedicel scar and strands extending into endocarp sinus), 4915, 5153, 7037, 7542, 8535, 8553, 8664, 12887, 12968, 13487, 17153, 20452, 20543–20544, 24310. Isolated endocarps 612, 12887, 18873.

Genus Cocculus DC. Cocculus lottii sp. n. (Pl. 32k–r) Diagnosis: Endocarp bisymmetric, dorsiventral length 9.2–9.6 mm, width 9–10.5 mm (n = 4), slightly laterally flattened, almost circular in lateral view with very broadly rounded dorsal surface, no dorsal crest but with a groove in the plane of bisymmetry; base convex to almost straight. Central depression small (up to 2.2 mm in diameter) reniform to subcircular, present on each lateral face but does not penetrate completely (i.e. not forming a foramen). Central condylar area surrounded by an incomplete annular bulged area which covers at least two thirds the distance from the central depression to the periphery. This is surrounded by a narrow, poorly developed groove separating it from a peripheral bulged ridge adjacent to the margin. Both lateral and peripheral bulges ornamented with irregularly arranged radiating fine and coarse rugulae, mostly transversely arranged, sometimes extending into short spines. Etymology: The specific epithet recognises Terry A. Lott for his contributions to palaeobotanical research at the University of Florida. Comments: These menispermaceous endocarps are distinguished from others in the Messel assemblage by being inflated in the plane of bisymmetry, and by the presence of a groove, rather than a keel, in the plane of bisymmetry. It is also noteworthy that they are nearly circular in lateral outline and that the condyle forms a central reniform depression on each of the lateral surfaces, with only a very narrow slit (sometimes not visible) extending to the attachment area. In these respects, and in overall morphology, they closely resemble endocarps of extant Cocculus. Circumferential bulges and grooves in combination with a reniform or lunate central depression are strikingly similar in e.g. Cocculus orbiculatus D. C. (Jacques 2009: fig. 51), but its rugulate surface does not extend into spines. 43


The ornamentation in Cocculus lottii varies from rounded radial ribs and/or tuberculae (pl. 32k, q) to more spiny tuberculae (pl. 32n, o), or even more finely ribbed without spines (pl. 32m). This may reflect relative maturity or the variable loss of mesocarp tissue in some, but not all specimens. Cocculus has distinctive endocarp morphology with variations in sculpture that are useful in distinguishing among extant species (Forman 1974). Hoot et al. (2009) produced a phylogenetic hypothesis based on sequence data in which extant Cocculus, as traditionally circumscribed, is nonmonophyletic: C. orbiculatus being more related to Hyperbaena Miers and Haematocarpus Miers than to the clade of C. balfouria Schweinf. ex Balf. and C. pendulus Diels. We did not see endocarps of C. balfouria, but those of C. pendulus (Jacques 2009: fig. 5k) are smoother and less similar to this fossil species than is C. orbiculatus. Other extant species, not included in the molecular study, that also have ornamentation similar to the C. lottii include C. hirsutus (L.) Diels (the type of the genus) and C. carolinus. We have also illustrated a smaller specimen, not included in the measurements of our diagnosis (SM.B Me 20562, pl. 32r). The specimen is about 2.8 mm thick, and 5.2–5.3 mm in diameter and retains a portion of pedicel. It may represent an immature or abortive specimen of C. lottii, but the possibility remains that it is a mature fruit of yet another taxon. Specimens: Holotype SM.B Me 8559 (here designated, pl. 32o). Paratypes SM.B Me 2228, 2516, 4152, 4242, 7177, 7187, 7214, 7718, 8502, 13030 (cluster), 20546, 20562, 21342.

Unnamed Menispermaceae (Pl. 34k) Description: Endocarp strongly curved, apparently nearly circular in the plane of bisymmetry. Ca. 5 mm wide across the plane of symmetry, and ca. 4 mm wide in the plane of symmetry. Specimen obliquely-dorsally compressed revealing three spiny ridges interpreted as a dorsal crest between a pair of lateral crests. Low narrow ridges radiating from the condylar projection toward lateral crest. Endocarp ornamented with up to five verrucae across area between lateral and dorsal crests. Comments: This specimen is unusual in being wider across the plane of bisymmetry than its width or height. The symmetry of the endocarp appears similar to that of extant Limacia scandens Lour., but that genus has a smooth, rather than spiny endocarp. As only one specimen is known, it appears to be in an unusual oblique compression, and internal structure has not been documented, we have not named it. Specimen: SM.B Me 4091.

44

Genus Tinomiscoidea Reid & Chandler Tinomiscoidea jacquesii sp. n. (Pl. 34a–c) Diagnosis: Endocarp shallow, boat shaped, obovate in face view, length 13–15.5 mm, width 8–9.5 mm, thickness not directly measurable, estimated c. 1–1.5 mm, bilaterally symmetrical, flattened dorsiventrally, apically pointed, basally rounded to rounded acute. Ventral side showing median longitudinal groove in the plane of bisymmetry. Ventral side showing transversely to slightly obliquely oriented striae diverging from the median line, dorsal side finely verrucate. Etymology: This species is named after F rédéric Jacques, recognising his contributions to the morphological and anatomical investigation of Menispermaceae. Comments: Although the wall thickness is not seen directly in these specimens, it is deduced to have been relatively thin because in the compressed condition, topographic features normally visible only from the ventral side can be seen from the dorsal surface. Morphology of these endocarps conforms to Tinomiscoidea Reid & Chandler 1933. Detailed comparison of these compressed endocarps with the two previously recognised species T. scaphiformis Reid & Chandler 1933 and T. occidentalis Manchester 1994 are precluded as those are locule casts with only fragments of the endocarp attached to the former. Tinomiscoidea is very similar to extant Tinomiscium according to Reid & Chandler 1933, but Tinomiscium is larger and has a shallower and smaller ventral concavity without a faceted margin. This species shows similarity with Tinomiscium petiolare Miers in surface sculpture and endocarp morphology (Jacques 2009: fig. 12l) but it is considerably smaller (length 15 vs 22 mm). Specimens: Holotype SM.B Me 2066 (here designated, pl. 34a). Paratypes SM.B Me 638, 2067, 2068, 13755, 16752.

Genus Parabaena Miers Parabaena cf. europaea Czeczott & Skirgiełło (Pl. 34d–f) Description: Endocarp boat shaped (only dorsal surface seen), elliptical in face view, length 8–9.2 mm, width 6.4–6.6 mm, bisymmetrical about a median dorsal ridge. Base rounded; apex also rounded but slightly more pointed than base. Surface coarsely transversely striate, striae interspersed with small nodules. Spines, up to 0.4 mm long, clearly evident protruding from periphery of compressed specimens. Comments: This species appears to be morphologically identical to Parabaena europaea Czeczott & Skirgiełło


(1967), recognised from the Miocene of Berzdorf, Saxony, Germany (Czaja 2003), in corresponding ornamentation, with very fine transverse ridges over the dorsal surface and the distribution of spines, but the endocarp of that species is somewhat smaller, 5.8–7.3 mm long. The shape of the two known Messel specimens is strongly similar to the modern Tinosporeae. Spiny boat-shaped endocarps occur in several extant genera of the tribe (e.g. Calycocarpum Nutt. ex Torr. & A. Gray: fig. 4G in Jacques 2009, ca. 16 mm long; Kolobopetalum Engl.: fig. 7E–G in Jacques 2009, 8–11 mm long; Parabaena Miers: fig 9D–G in Jacques 2009, fig. 1C–L in Forman 1984, 6.6–9 mm long; Rhigiocarya Miers: fig. 10C in Jacques 2009, 14 mm long; Tinospora: 11.6–7, 11.17–18, 5–7 mm long). The endocarps of some of these genera are much larger than the Messel specimen whilst those of Tinospora have only scant tiny spines and thus also differ from the Messel specimen. A further comprehensive survey is necessary to recognise the modern affinity. The species Tinospora hardmanae Manchester (1994) from Clarno is also known to have been spiny, but is smaller than the Messel specimen. The affinity of the Clarno species to the extant genus may be questionable because we are not aware of spines in the extant Tinospora as recently figured by Jacques (2009). Tinospora wilkinsoni Chandler 1961b is only slightly less elongate than the Messel specimen and is 8 mm long. It has short conical spines with most of the tips broken during fossilization and is therefore rather similar to the Messel specimen but lacks the transverse striation. T. excavata Reid & Chandler emend. Chandler 1961b from the London Clay is smaller (less than 7 mm long) and is ornamented with tubercles and so is clearly distinct from the Messel specimen.

tions, one showing the ventral surface and the other the dorsal surface, so that we have not confirmed their conspecifity by comparing like surfaces. The boat-shaped morphology corresponds to that common to endocarps of Tribes Tinosporeae and Tinomisceae. In size and ornamentation, these specimens are rather similar to Tinospora rugosa Reid & Chandler. However the London Clay species shows a marginal ridge at the junction of dorsal and ventral surfaces, that we have been unable to confirm, has a less pronounced median ridge, and the apex is not pointed. These differences may be partially due to the different preservational modes, because the pyritized London Clay specimen retains the three dimensionality of the original endocarp, while the Messel specimens have been compacted and likely somewhat distorted. The two specimens are similar in form and sculpture to Sarcolophium suberosum (Diels) Troupin, as figured by Jacques (2009: fig.10D), but detailed comparisons have not been made. For comparison with other fossil Tinosporeae see under Parabaena above. Specimens: SM.B Me 2216, 2218.

Tinosporeae sp. 2 (Pl. 34i, j) Description: Endocarp boat-shaped, strongly dorsiventrally flattened, slightly concave ventrally and correspondingly convex dorsally. Ovate in face view, rounded basally, obtusely pointed and rounded apically; 6.5 mm long, 5 mm wide, c. 1 mm thick; with a faint, straight median ridge along the plane of bisymmetry, more prominent on the ventral side. Surface smooth.

Specimens: SM.B Me 4023, 13741.

Tribe Tinosporeae Hook. & Thomson Tinosporeae sp. 1 (Pl. 34g, h) Description: Endocarp ovate in face view, 4.5–6.5 mm long, 4–5 mm wide, thickness 0.9 mm, slightly more pointed at one end, dorsiventrally flattened. Bisymmetrical about median ridge visible on both dorsal and ventral surfaces. Ventral side smooth, shallowly concave with narrow median longitudinal ridge. A fibrous strand runs along the crest of the ventral median ridge. Dorsal side convex, verrucate to irregularly rugulate with a thick median longitudinal ridge that becomes more raised at the more pointed end of the fruit. Outer layer thin, very finely rugulate/verrucate and readily detaching from the smooth inner layer. Entire surface clothed in hairs. Comments: The two specimens cited below are believed to represent the same species, based on similar size and shape. However, they are preserved in different orienta-

Comments: This endocarp is similar in shape and general organization to Tinosporeae sp. 1, but lacks ornamentation. Specimen: SM.B Me 2213.

Tinosporeae sp. 3 (Pl. 34l–n) Description: Endocarp very rounded boat-shaped, almost circular in lateral profile but with a flattened ventral surface, with a pronounced ventral cavity. Elliptical in face view, with a very slight terminal angle. Length in the plane of bisymmetry 8.3–8.5 mm, width of the ventral face in ventral view (strongly distorted by plane of compaction) 4–7.5 mm (n = 2). A narrow sharp keel extends apically from the ventral cavity over the apex and all around the dorsal face bisecting the specimen in the plane of bisymmetry. Placental scar not observed. Surface ornamented by distinct discrete rounded tubercles (0.7–1 mm diameter) protruding from the surface less than 0.1 mm. 45


Comments: Specimens of this taxon might be confused with Tapiscia if the plane of compaction does not reveal details of the keel or the ventral cavity. However, this species has a thin median dorsal ridge in the plane of bisymmetry and has larger, more pronounced and clearly discrete tubercles in contrast to Tapiscia which lacks the median dorsal ridge and has finer verrucae or rugulae. The combination of the ventral concavity, bilateral symmetry and keel are features characterizing extant genera of the Tinosporeae tribe of the Menispermaceae. In particular, the genus Tinospora frequently shows similar endocarp morphology, and according to Chandler (1961b) many living species of Tinospora have tubercled endocarps. The Messel material is very similar to T. excavata Reid & Chandler emend Chandler 1961b from the London Clay, although the specimens from Messel are slightly larger and have a larger ventral concavity. The extant genus Tinospora as traditionally circumscribed appears to be polyphyletic (Jacques et al. 2011), and more work is needed to determine the relationships among extant and fossil taxa with this kind of endocarp morphology. Specimens: SM.B Me 2187, 2190, 2193, 8830, 16704.

Family Myristicaceae R. Br. Genus Myristicacarpum Gregor Myristicacarpum sp. (Pl. 34o, p) Description: Seed circular in profile view, 8 mm high, 9 mm wide, apex and base rounded, lenticular in cross section (crushed). Seed surface smooth. Transverse fracture section showing ruminate endosperm with irregular longitudinal plate-like intrusions of the seed coat c. 0.15–0.2 mm thick that extend radially inward toward centre of the seed, where some of them merge to form V shapes as viewed transversely. Comments: Myristicacarpum Gregor (1977) is used to accommodate fruits and seeds with ruminate endosperm comparable to that of extant Myristicaceae. The only other known Paleogene occurrence is from the London Clay flora (Doyle et al. 2008). The seed from Messel is shorter and more globose than the single specimen of M. chandlerae Manchester, Doyle & Sauquet. Specimen: SM.B Me 5605.

?Nymphaeales Salisb. ex Bercht. & J. Presl Unnamed seeds (Pl. 35a–i) Description: Seeds either isolated (SM.B Me 4045), in groups of few seeds, or in closely packed clusters of 100 or more. Seeds ovoid, 3–4 mm long and 2–3 mm 46

wide (length/width ratio about 1.33), anatropous with a sub-basal chalaza (seen as a round protrusion or a scar of damaged tissues in most specimens, e.g. pl. 35d) and a terminal circular apical cap (embryotega – operculum) carrying a slit-like micropyle and slightly protruding circular hilum (pl. 35d, f, g). The micropylar slit crosses the margin of the hilar scar. Longitudinal ridge present on many seeds and although we have not observed vascular tissue we interpret this ridge as a raphe ridge running between the apical hilum and the basal chalaza. Seed surface minutely papillate and pitted. Seed coat consisting of a cuticular inner layer and an outer sclerotic layer. Surface of outer layer showing interlocking digitate cell outlines (pl. 35h) with micropapillate to micropitted surface ornament (pl. 35i) often obscured by additional surface coating. Cell outlines irregularly but deeply lobed with 8–11 lobes per cell. Most lobes are single, but occasional irregular bifurcations occur. Cells not arranged in regular longitudinal rows but irregularly distributed over the surface. Polygonal cells on the cap. Seed coat thickness cannot be determined due to compression. Furthermore anatomy of the seed coat in sections is obscured, even with SEM and TEM. Tegmen cuticle (but not micopylar detail) has been observed by epifluorescence microscopy, showing characteristic convoluted anticlinal cell walls in conical area beneath the micropyle (observed in isolated seeds and seed clusters). The cuticles have a pattern of oblong cell outlines with long axis orientated perpendicular to the long axis of the seed. Comments: These seeds conform to the Nymphaeales in their anatropous derivation (deduced from longitudinal ridges interpreted to contain the raphe), presence of operculum (e.g. pl. 35d, g) and by the interlocking undulate cells of the surface (pl. 35h). The massive clusters of seeds sometimes encountered (e.g. pl. 35a, b) may be an indication of being held together by slime as in the case of extant Nuphar Sm. The seeds are not named here as they are part of a separate ongoing systematic study (Collinson, Wilde and Borsch, unpublished data) on the seeds and the large flowers in which the corresponding ovules occur. Seeds are known to occur in mature flowers (e.g. SM.B Me 14122, 7012a (showing stamens), 1712) of the kind figured by Schaarschmidt (1986). There remains a possibility that this taxon does not belong in Nymphaeales but represents an extinct clade within the “ANITA” group, hence we place a question mark in the ordinal assignment. Specimens: Seed clusters: SM.B Me 2735, 7011, 7014. Seeds with the operculum in place: SM.B Me 2169, 4057 (two seeds together), 4483, 4484, 4627, 4614, 4762, 4798, 4900, 5092–5094, 7260, 7618, 7627, 7666, 7693. Seeds without the operculum: SM.B Me 2167, 2619, 4625, 4673, 4750, 5095, 7333, 8206, 8236. Other specimens: SM.B Me 21625. The following specimens/fragments of specimens were examined by SEM (isolated specimen SM.B Me 4045, clusters 7017a and 2735, and flower 7012b). In addition, SM.B Me 7012b from a seed cluster, and 2736a from a flower were examined by TEM.


Family Nyssaceae Juss. ex Dumort. Genus Nyssa L. Nyssa disseminata (R. Ludw.) Kirchheimer (Pl. 36a–d) Basionym: 1857 Pinus disseminata Ludwig, Palaeontographica, 5: 89, pl. 20, fig. 2 Synonymy: 1937 Nyssa disseminata (R. Ludwig), Kirchheimer, Braunkohle, 36: 916, fig. 11.

Description: Endocarp ovoid to ellipsoid, length 12– 14 mm, width 7–9 mm (n = 4), base and apex rounded to slightly pointed, smooth with 4–5 straight longitudinal furrows on the exposed surface (indicating a total of approximately 10). An arched slit representing the margin of the germination valve confined to the apical half of the fruit (as a phenomenon of compaction we can only see one germination valve). Germination valve 4.5–7 mm long, 4.8–5.7 mm wide. Comments: The presence of a well defined apically opening germination valve is characteristic of nearly all Cornalean genera. The germination valves of Nyssaceae including Nyssa and Camptotheca Decne. and the extinct genus Amersinia Manchester, Crane & Golovneva (Manchester et al. 1999) are confined to the apical half of the endocarp (Eyde 1963). The relative size of the germination valve and a smaller length/width-ratio are consistent with Nyssa for the Messel specimens. Eyde (1997) reviewed the morphology of extant and fossil Nyssa species. In the European Tertiary, Nyssa stones with broad ribs with intervening sunken bundles occur in the unilocular fruits, N. oviformis E. Reid (1930), silicified fruits from Brittany, France, presumably mid-Eocene. They are distinguished, however, by their larger size (15.5 × 9 mm) and apparently thicker wall. Nyssa disseminata (R. Ludw.) Kirchheimer, known from the Oligocene through the Pliocene includes stones most similar to those of extant N. sylvatica (Eyde 1997). Mai (1965, 1973) presented a narrower concept for this species than Kirchheimer (1937) such that N. disseminata now includes only stones with an oval or elliptic outline and one, rarely two, seed chambers. Germination valves are broad and more or less rounded, and the ribs are rounded. Size ranges from 8–14 mm long and 4–7 mm wide (Mai & Walther 1978). Nyssa ornithobroma Unger is distinguished from N. disseminata by a longer and more pointed germination valve than N. disseminata. Typically, N. ornithobroma stones are obovate – broadest above the midline and the number of locules varies from one to four, even in the same population. The emended diagnosis (Mai & Gregor 1982) gives N. ornithobroma's size as 7–22 × 4–12 mm. Nyssa boveyana Chandler (1957) has up to three locules and sharper ribs, some of which have vascular bundles running along their crests, rather than in the furrows, a feature similar to extant N. aquatica L. (Eyde 1997). Specimens: SM.B Me 7194, 7201, 7207, 7566, 8256.

Family Pentaphylacaceae Engl. This family includes many taxa formerly placed in Theaceae (APG III 2009).

Genus Cleyera Thunberg Cleyera sp. (Pl. 36e–g) Description: Seed campylotropous, laterally flattened, rounded in outline, truncated at micropyle and chalazal end, shallow median depression (inferred condyle) around which the C-shaped embryo cavity curves. Width 4 mm measured across limbs of the “C”, height 3.3 mm. Surface finely reticulate. Testa surface with a polygonal cell pattern, cells with upraised outer periclinal walls, arranged in irregular, more or less concentric rows (about 13 cells/mm), following the curvature of the margins of the lateral face of the seed. Cells more elongate over the central area. Comments: In campylotropous shape and surface sculpture these seeds resemble those of the extant genera, Cleyera and Eurya Thunb. of the theaceous subfamily Ternstroemoideae, now treated as Pentaphylacaceae (APG III 2009). In the identification of this genus from the Eocene of Oregon, Manchester (1994) followed Friis (1985) who argued that seeds of Cleyera tend to be more strongly campylotropous and usually larger than those of Eurya. The Messel seeds are large, relative to other fossil species, which are usually less than 2.5 mm in maximum dimension. Otherwise, these specimens closely resemble those that have been assigned to Eurya stigmosa (Ludwig) Mai from the Paleocene to Upper Miocene of west and middle Europe (Mai 1971, Mai & Walther 1978, 1991). Specimens: SM.B Me 4914, 8087, 8111, 8897, 12164, 17394, 20158.

Family Rhamnaceae Juss. Genus Berchemia Neck. ex DC. Berchemia mellerae sp. n. (Pl. 36h–n) Diagnosis: Endocarp spathulate in face view, length: 7.8–11.0 (average 9.7) mm, width 5.8–7.2 (average 6.5) mm (n = 5), thickness in the plane of compression 1.2 mm, rounded apically, sharply truncate basally, smoothly rounded lateral margins. Truncated end with a pair of very shallow to indistinct concavities separated by a median ridge. Surface smooth, with a broad median ridge which is continuous, with a distinct narrow median groove running right around the entire specimens. One specimen retains a portion of apparent outer softer tissue 1.5 mm thick at the base and left side. 47


Etymology: This species is named in honor of Barbara Meller in recognition of her contributions to Tertiary palaeocarpology. Comments: The two planes of symmetry with the median ridge and groove running around the entire specimen with a prominent truncation/cavity at the base of the endocarp corresponds precisely to the condition in endocarps of extant Berchemia (e.g. pl. 36o). The only other fossil fruit of this genus of which we are aware is from the Kenyan Miocene named Berchemia discolor Chesters 1957. Being a mineralized cast it is difficult to compare with the Messel material. However, Chesters (1957) noted the bisymmetric endocarp with septum between two one-seeded locules marked internally by a shallow groove; distal end rounded and proximal end truncated with oval apertures. All of these features conform to our specimens (allowing for preservational differences) and the endocarp sizes are similar (12 mm × 7 mm for the African material). However, Chesters (1957) described and figured (pl. 20, fig.1) a rough rugose layer of exocarp and a nodular outer layer of endocarp whereas exocarp and endocarp are smooth in our material. Therefore, we recognise our material as a new species, considerably pre-dating the only other fossil occurrence known to us. The Rhamnaceae are diverse in fruit morphology. Among those with drupes such as Ziziphus Mill., Berchemia, Rhamnidium Reissek, there are intergeneric differences. Ziziphus has thick woody endocarps with ribbed sculpture and lacking obvious straight longitudinal grooves. Rhamnidium (based on observation of R. elaeocarpum Reissek) has endocarps that are similar in symmetry to this fossil with median groove on both faces, but lack the basal cavity/truncation. Karwinskia Zucc. has ellipsoidal endocarps, lacking the basal cavity and having the grooves running along the edges of the endocarp rather than medially. To our knowledge, the features seen in this fossil are diagnostic for the genus Berchemia. We compared with B. floribunda (Wall.) Brongn. (A: J. F. Maxell 90–267, from Thailand), and B. lineata DC. (A: E. H. Wilson 11800 from Taiwan, pl. 36o). The endocarps are lensoidal in cross section, and relatively thin walled, so it is not surprising that the fossils are collapsed/flattened in the wider plane of symmetry. The apparent adhering portion of fleshy outer tissue seen in one of the fossils is consistent with a drupaceous fruit of Berchemia. Specimens: Holotype SM.B Me 4184; Paratypes SM.B Me 4706, 4773, 5585, 7113, 7366, 8793, 10597, 18071, 19004, 19183, 19445, 21298.

Family Rutaceae Juss. The Rutaceae, now placed within the order Sapindales, contain several clades of which the Citrus L. clade (which now includes Ruta L. and should be referred to as Ruteae; 48

Scott et al. 2000), Ptelea L., and the Zanthoxylon Walter clades are most familiar. According to molecular phylogenetic analyses, Toddalioideae are a clade that includes Zanthoxylum, Phellodendron Rupr., Toddalia and Tetradium Dulac, which is, in turn sister to a clade that includes Dinosperma T. G. Hartley, Bosistoa F. Muell., Halfordia F. Muell., Euodia Gaertn., Melicope J. R. Forst. & G. Forst., and Acronychia Forst. (see Poon et al. 2007). Collinson & Gregor (1988) discussed issues of fossil seed nomenclature and concluded that the name Rutaspermum was appropriate for fossil seeds from Messel which share some characteristics with Zanthoxylum, Toddalia and Acronychia.

Genus Rutaspermum Chandler emend. Gregor Rutaspermum chandleri Collinson & Gregor (Pl. 37m) 1988 Rutaspermum chandleri Collinson & Gregor, Tertiary Research, 9: 68–70, pl. 1, figs 1–7, pl. 2, figs 1–3, pl. 3, figs 3, 4, 9–12

Comments: This species has a smooth-surfaced seed. There are no new additional specimens from the oil shale which means that this species is almost entirely restricted to site SMF 7 with only SM.B. Me 4132 (pl. 37m) and one other specimen tentatively assigned to the species from the oil shale. See R. messelense below for further comments. Specimens: Those of Collinson & Gregor (1988).

Rutaspermum messelense Collinson & Gregor (Pl. 37a–e) 1988 Rutaspermum messelense Collinson & Gregor, Tertiary Research, 9: 70–71, pl. 1, figs 8, 9, 15–19.

Comments: R. chandleri is a smooth-surfaced Rutaspermum seed whereas R. messelense has a pronounced surface reticulum of ridges and depressions on which are superimposed fine pits. The best preserved specimens of the latter species have a thin outer testa layer of pentagonal cells which hides the fine pitting, and giving the seeds a smooth, shiny, reticulate appearance. Some R. messelense have much less pronounced reticulum than others and may almost appear smooth unless studied carefully. Some specimens have more pronounced longitudinal ridges with subordinate transverse ridges whilst others have equally developed ridges. There is a continuum of variation among the morphologies within R. messelense. The most similar fossil species is Acronychia ornata (Chandler) Mai from the Geiseltal and Hordle (Mai 1970) but these are larger and have more pronounced longitudinal ribs than R. messelense. In contrast to R. chandlerae, R. messelense is abundant in the oil shale and very rare at site SMF 7. This suggested that the latter species grew


around the Messel lake whereas the former did not (Collinson & Gregor 1988) a conclusion substantiated after twenty additional years of collecting and research on the oil shale. Specimens: Holotype SM.B Me 2362 (Collinson & Gregor 1988: pl. 1, fig. 8, 9). Additional specimens include those of Collinson & Gregor (1988), plus SM.B Me 2587, 4613, 4818, 5071, 5069, 5070, 7124– 7126, 7295 (with boring), 7739, 7945–7952, 7953 (shows dehiscence), 7954–7962, 7963 (shows dehiscence), 8254, 8302, 8342, 8422, 8514, 8530, 8557, 8590, 8654, 17137.

eral faces of seed convex with no obvious ornamentation, but surface finely pitted with a pattern of polygonal cells. Comments: This seed type is similar to Rutaspermum chandlerae in the smooth surface but R. chandlerae has a flat hilar face and a smooth area around the hilar scar. One specimen (SM.B Me 4142, pl. 37j) shows a slit which may be part of a dehiscence slit. The slit would bisect the specimen which is unlike R. chandlerae and like Toddalia (Collinson & Gregor 1988). Specimens: SM.B Me 4142, 4677, 4304.

Rutaceae undetermined A sensu Collinson & Gregor (Pl. 37f) 1988 Rutaceae undetermined A Collinson & Gregor, Tertiary Research, 9: 71, pl. 1, figs 10–12.

Comments: Undetermined A is a typical elongate boatshaped Rutaceae seed with flattened hilar scar, raphal excrescence and micropylar knob. The surface ornamentation is dominated by longitudinal ridges with rare anastomoses. There are no additional specimens to give further indication towards the systematic position within the family. Specimens: Those of Collinson & Gregor (1988) and SM.B Me 4037.

Rutaceae undetermined B sensu Collinson & Gregor (Pl. 37g–i) 1988 Rutaceae undetermined B Collinson & Gregor, Tertiary Research, 9: 71, pl. 1, figs 13–14.

Comments: Undetermined B is a laterally flattened seed hence ellipsoidal in lateral view. It conforms generally to seeds of Rutaceae but critical diagnostic characters are lacking. This seed is also ornamented with a faint reticulum but the elliptical shape distinguishes it from R. messelense. One possible additional specimen has been recognised. Specimens: Those of Collinson & Gregor (1988); SM.B Me 7372 (pl. 37g, h) may be another specimen of this taxon.

Genus Toddalia Juss. cf. Toddalia sp. (Pl. 37j–l) Description: Seed ellipsoidal, length 4.8–6.0 mm, lateral thickness 4.5 mm, dorsiventral thickness 4.2 mm, slightly flattened on the hilar surface, with an elongate-elliptical hilar scar almost terminal on the ventral face. Hilar scar raised, not flattened, not surrounded by a smooth area, and occupying one third to one half the length of the ventral face. No prominent raphal excrescence, micropyle not evident. Lat-

Family Sabiaceae Blume Genus Meliosma Blume Meliosma sp. (Pl. 38a–f) Description: Endocarp subglobose, bilaterally symmetrical (length in plane of symmetry 11.5 mm, width across plane of symmetry 10.5 mm, dorsiventral height 11 mm), with a keel in the plane of symmetry and a ventral depression with a prominent funicular cavity or plug (SM.B Me 10620, pl. 38a, b). Surface smooth to finely scabrate, but not shiny. Comments: The characters clearly place these endocarps in Meliosma (van Beusekom 1971, Collinson 1983a). This species is apparently rare at Messel although specimens with different planes of compression may have been overlooked. In size, which is large in relation to most European fossil representatives of this genus, as well as morphology, this species conforms well to Meliosma jenkinsii Reid & Chandler (1933) but detailed anatomical comparisons of the London Clay type material with these compression fossils has not been done. Specimens: SM.B Me 8818, 10620, 21304. A similarly compressed smaller specimen, 5 mm in diameter, may represent another species of Meliosma (SM.B Me 4111, pl. 38g–i).

Family Sapotaceae Juss. Genus Sapotispermum Reid & Chandler cf. Sapotispermum sp. (Pl. 38j, k) Description: Seed elliptical, length 18 mm, width 11.5 mm. In lateral view with a strongly oblique truncation, 10 mm long, about 30° angle to the long axis, bilaterally symmetrical, surface smooth. Traces of thin dark cuticle remain on the surface. Comments: Large elliptical bilaterally symmetrical seeds with a pronounced oblique facet are characteristic for seeds of Sapotaceae. Sapotispermum sheppeyense Reid & Chandler is obpyriform and smaller than the Messel specimen (Collinson 1983a). 49



Family Simaroubaceae DC. Genus Ailanthus Desf. Ailanthus confucii Unger (Pl. 38l, m) 1850 Ailanthus confucii Unger, Denkschr. Kaiserl. Akad. Wiss Wien, Math.-Naturwiss. Cl., 1: 23 [1850c]. 1859 Heer (first picture of Unger’s original specimen), Flora tertiaria Helvetiae. Vol 3. J. Wurster, Winterthur 87, pl. 127, fig. 36.

Further synonymy provided by Corbett & Manchester (2004). Description: Fruit biwinged, elongate elliptical, tapered at both base and apex, with a central seed. Fruit 14, 17.5, 24.5 mm long, 3, 3.7, 5 mm wide, seed elliptical, 2–4 mm in diameter. Fine, subparallel veins extending longitudinally. For detailed description see Corbett & Manchester (2004). Comments: The Messel specimens belong to a widespread morphospecies which is also known from North America and Asia (Corbett & Manchester 2004). Although the majority of specimens in Messel are relatively small in size (less than 15 mm in length) compared to most specimens from other floras, a single specimen (SM.B Me 21808, pl. 38m) with a length of 25 mm indicates that the population from Messel is consistent with the range of dimensions normally covered by the species. Comparisons with the extant species by Corbett & Manchester (2004) indicate the closest similarity with A. altissima of China. Specimens: SM.B Me 4006, 4232, 4233, 4747, 4785, 4786, 16837, 21808, 23395, 24010.

Family Tapisciaceae Takht. Genus Tapiscia Oliv. Tapiscia pusilla (Reid & Chandler) Mai (Pl 39a–l) Basionym: 1933 Palmospermum pusillum Reid & Chandler, London Clay Flora 115, pl. 1, fig. 32–34. Synonymy: 1976 Tapiscia pusilla (Reid & Chandler) Mai, Abh. Zentr. Geol. Inst., 26: 122

Description: Fruit subglobose to pyriform, length 5.8– 8.5 mm, width 5.0–7.0 mm, diameter 1.0–1.5 mm, bilaterally symmetrical, rounded distally, the opposite end tapering (marking the pedicel of the fruit and the micropyle of the enclosed seed). Dorsal side rounded, ventral side with a prominent obovate to triangular concavity (representing the chalazal scar beneath). Surface of pericarp with small regularly spaced scabrae, verrucae or short rugulae. Longitudinal strands linking between the ventral 50

depression and the micropylar protrusion (representing the underlying position of the raphe). Comments: These specimens conform to Tapiscia based on size and the characteristic ornamentation corresponding precisely to the single extant species, T. sinensis Oliv. The presence of this genus in the European Tertiary was first recognised by Mai (1976, 1980). Relatively few Messel specimens are oriented in a plane revealing the critical diagnostic characters of Tapiscia which are visible only on the ventral surface (e.g. pl. 39a–d, g). However, SRXTM was applied to a specimen still buried in shale with only its dorsal surface exposed (pl. 39i). The distinctive chalazal depression, micropylar protrusion, and thin pericarp over a smooth seed, are clearly seen in the resulting digital sections (pl. 39j–l). Another specimen was physically removed from the shale to reveal both ventral and dorsal surfaces (pl. 39g, h). The Messel specimens correspond in morphology and ornamentation to the species T. pusilla (Reid & Chandler) Mai from the London Clay. Seed moulds that could represent the same species were described as T. subglobosa Mai (Mai 1976, 1980) from the Middle Eocene of the Geiseltal, however as the external characters of the Geiseltal specimens are unknown, we hesitate to combine the species. The North American Middle Eocene species T. occidentalis Manchester (Manchester 1988, 1994) is virtually identical in morphology to this species, differing only by its smaller size. Although preserved as compressions, Messel specimens are not dried and therefore are inferred to have retained their original size. Some specimens show large aggregations of these seeds (SM.B Me 8735; 24289, pl. 39e), without intervening tissue, or pedicels, suggesting coprolite association. One of the fruit specimens (SM.B Me 17741) appears to be attached to a very swollen peduncle (cuticle preserved and folded), but no perianth scar is present (pl. 39f). If this represents a pedicel it is more inflated than in the extant species. Specimens: SM.B Me 2189, 2246, 2276, 2530, 4167, 4176, 4191, 4194, 4431, 4601, 8735, 12234, 12243, 13060, 13876, 14862, 14875, 14833, 16874, 17741, 18136, 18142, 18144, 19502, 21386, 24289. Other specimens are likely to exist in the collections but, as explained above, if the ventral surface is not exposed the ornamentation and size can only be taken as tentative indications of affinity.

Family Theaceae Mirb. Genus Camelliacarpoidea gen. n. Camelliacarpoidea messelensis sp. n. (Pl. 40a–i) Diagnosis: Fruit subglobose, fruit body height and width 16.5 mm, as preserved, finely rugulate, apparently leathery, flattened by compression. Apex missing, stylar condition unknown. Fruit containing at least two brittle, cracked, crushed elongate seeds, with a shiny seed coat.


Persistent hypogynous calyx with four clearly seen imbricate lobes, (5 or 6 total inferred from symmetry), with greatly thickened circular basal disk, 6 mm diameter, below the lobes, peduncle 1.8 mm thick, 2.9 mm long; calyx lobes 4 mm long (measured from edge of disk), 3.5 mm wide. Calyx and disk with a peculiar strongly verrucate and rough texture. Each calyx lobe with a median adaxial faint keel incomplete/obscured, placentation and morphology unknown. One seed measurable: 4.3 mm long and 3.4 mm wide. Seed with a uniseriate columnar sclerotesta 200 µm thick, with polygonal cell outlines in surface view 20–30 µm in diameter, each polygon with an indistinct central pit. Sclereids thick walled with small lumina, ca. 2–3 µm, decreasing in size outwards (pl. 40i). Etymology: The generic name refers to the similarity of fruit morphology to that of extant Camellia L., and the specific epithet to the site of Messel. Comments: Fruits with hypogynous calyx of imbricate sepals, each with a median fold, and prominent disk are consistent with Theales (now treated as a part of Ericales sensu APG III 2009) and indicative of relationship to the Theaceae s.l. It is now fashionable to treat Theaceae as multiple families, including Pentaphylaceae (including Ternstroemioideae), and Theaceae s.str. as well as Sladeniaceae (APG III 2009).This fossil was probably a capsule as inferred from the longitudinal grooves, but there is no indication of a central column in the fractured fruit. It is similar in general organization, globose fruit, sepal configuration, and several seeds to Camellia (and Pentaphylax Gardner & Champ.?) but we have not examined the seed anatomy of these extant taxa for detailed comparison with the fossil. Specimen: Holotype SM.B Me 8953 (here designated, pl. 40a–i). No other specimens known.

Family Toricelliaceae Hu Genus Toricellia DC. Toricellia bonesii (Manchester) Manchester (Pl. 39m–p) Basionym: 1994 Tripartisemen bonesii Manchester, Palaeontographica Americana, p. 113, pl. 62, figs 8–15. Synonymy: 1999 Toricellia bonesii (Manchester) Manchester, Ann. Missouri Bot. Gard. 86: 522 fig. 1c–f.

Description: Endocarps 2.5–4.5 mm in maximum dimension (n = 5), originally probably globose to subglobose, now strongly compressed in various planes. In dorsi-ventral compression (pl. 39m) the centre of the ventral surface has two prominent ridges which are merged at their ends as seen at the endocarp margin and diverge up to 1.5 mm apart as they cross the endocarp. In

lateral compression these ridges are seen to pass around two-thirds of the circumference of the endocarp and demarcate a curved seed chamber (SM.B Me 2197, 2198, 5745). At one end of the seed chamber a transverse ridge crosses the space between the main ridges joining them and marks the base of a sub-triangular germination valve up to 1.9 mm long and 1.5 mm broad. A third median ridge may or may not be present along the full extent of the seed cavity but is always present on the germination valve (n = 2). Dorsal endocarp surface with two large holes either side of a median septum, holes occupying about two-thirds of each lateral-dorsal area. Each hole overlies a lateral endocarp chamber. In lateral compression only one hole is visible (e.g. SM.B Me 2197, 2198). SM.B Me 5745, which has been released from the oil shale and, although compressed, can be viewed from all angles, is a dorsi-lateral compression which clearly shows one lateral-dorsal hole and a median dorsal septum within the endocarp. Endocarp wall is thin (less than 0.2 mm). Cellular detail is not visible by LM. Comments: Most specimens compressed in the oil shale reveal few diagnostic characters and are difficult to recognise (e.g. pl. 39n). Sometimes lateral compression (D-shaped profile) shows, on the curved edge, two concentric ridges with part of the germination valve at one end of the seed chamber (pl. 39o). The holes in the chambers on the dorsal face, seen in lateral view in pl. 39o, p and in lateral-dorsal view on SM.B Me 5745, are easily mistaken for damage. As a consequence, Collinson (1988, table 1 caption) only recognised a single specimen from the oil shale, SM.B Me 5745. Using more abundant specimens from site SMF 7 Collinson (1988, 1989a) tentatively suggested an affinity with Lythraceae interpreting the specimens as seeds and the holes as bite marks. Meller (in Meller & Collinson 1999) and Meller (2006) recognised the correct systematic affinity as endocarps of Toricellia. New specimens, especially SM.B Me 12260 (pl. 39m), displaying the complete seed chamber and germination valve, have enabled this to be confirmed herein for the oil shale specimens. Meller (2006: 319) refrained from assigning her Miocene specimens from Oberdorf to the species T. bonesii (Manchester) Manchester (1999) with certainty on the basis of the fact that it was not possible to unequivocally confirm the presence of the dorsal endocarp holes in the type assemblage from the Eocene of Clarno. Subsequently, we have sectioned additional specimens from the Clarno Nut Beds locality (e.g. UF225-9290), and confirmed the presence of holes in the lateral walls of the lateral chambers. Meller also noticed a size difference between the three assemblages. M eller (2006) quoted C ollinson (1989a) as giving a size range of 1–2 mm for the Messel specimens but measurement was based exclusively on specimens from site SMF 7. The Messel oil shale specimens, the Clarno specimens and the Oberdorf specimens all overlap in 51


size. This is the oldest known European record for the genus, which is also known from the Paleocene of North Dakota and Middle Eocene of Oregon and Washington, in western North America (Manchester et al. 2009). Specimens: SM.B Me 2197, 2198, 4622, 4812, 5745 (loose from oil shale) 8156, 8157, 8352, 8452, 12260.

Family Ulmaceae Mirb. Genus Cedrelospermum Saporta emend. Manchester Cedrelospermum leptospermum (Ettingshausen) Manchester emend. Wilde & Manchester (Pl. 41a–f) Basionym: 1853 Embothrites leptospermus Ettingshausen, Abh. Geol. Reichsanst. Wien, 2: 51, pl. 14, figs 15–25. Synonymy: 1987 Cedrelospermum leptospermum (Ettingshausen) Manchester, Rev. Palaeobot. Palyn. 52: 122–124, pl. 1, 1–4; 1989b, Manchester, Amer. J. Bot. 76, fig. 45; Manchester 2003 Wilde & Manchester, Courier Forschungsinstitut Senckenberg, 241:147–153.

Description: Fruit a samara, 5–8.5 mm long and 2.8– 3.2 mm wide, with a flattened elliptical endocarp and a single oblique lateral wing. Veins of the wing, including marginal veins, of a single order converging distally toward V-shaped stigmatic notch at the distal end of the wing. Fruits borne in groups at alternate nodes on zig-zag twig. Comments: The species from Messel was already treated by Wilde & Manchester (2003). Additional new specimens have been acquired more recently, including a more complete leafy shoot with fruits attached (pl. 41b, d), and a leafless twig with several attached fruits (pl. 41c) supplements the one previously described. Cedrelospermum fruits are very rare in the Messel collections, represented only by those attached to the two leafy shoots and by a single isolated fruit. The leaves belonging to it, corresponding to the foliage morphogenus Tremophyllum Rüffle are common at Messel (Wilde 1989, Wilde & Manchester 2003), and correspond well with the leaves found attached to twigs bearing Cedrelospermum fruits from the Green River and Florissant floras of North America (Manchester 1989a). Specimens: SM.B Me 7274, 15393, 15974; an additional unnumbered specimen photographed in the year 2000 (pl. 41b, d) has not been relocated.

Family Vitaceae Juss. The grape family is well represented in the Messel fruit and seed flora, including both extant and extinct genera. Seeds of extant Vitaceae are readily recognisable by the presence of paired ventral infolds and rounded dorsal chalaza; subgroupings corresponding to extant clades 52

can be discerned by differences including ventral infold configuration, chalazal shape, presence or absence of apical grooves, and ornamentation (Chen & Manchester 2007, 2011).

Genus Vitis L. Vitis messelensis sp. n. (Pl. 42a–h, l) Diagnosis: Seed oval to pyriform in dorsal and ventral view, 4.8–5.3 mm long, 4.0–4.3 mm wide, dorsiventral thickness 1.5–1.9 mm as measured but original thickness uncertain because of compression of the fossils; some specimens are compacted laterally suggesting some degree of inflation in life. Apical notch shallow to missing. Beak prominent, cylindrical, truncate, protruding 0.2 mm. Ventral surface smooth. Ventral infolds narrow, nearly parallel near the base and more diverging apically, relatively short, about one-third to half of the full seed length. Raphe ridge sometimes with a slit-like groove. Dorsal surface usually smooth, but sometimes with very slight fluting radiating from the chalaza. Chalaza pyriform to elliptical, more or less centrally positioned (if offset slightly apical), approximately one third the length of the full seed. Chalaza-apex groove prominent, chalazabase groove less pronounced. Seed coat c. 250 µm thick, composed of a biseriate layer of anticlinally elongate cells. Etymology: The specific epithet refers to the site of Messel. Comments: Among extant seeds of Vitaceae, the combination of characters in this species is well accommodated in the genus Vitis. The smooth surface, pronounced truncate beak, elliptical chalaza, small, slightly divergent ventral infolds, and only slight apical groove, are well matched by the extant Chinese species, V. wilsoniae H.J.Veitch (fig. 16A in Chen & Manchester 2011). Vitis messelensis seems to be the most common vitaceous seed type in the Messel collections. In addition to the numerous isolated seeds, a fruit containing two seeds is assigned to this species (pl. 42l), and another fruit with one seed exposed in dorsal view may also belong (SM. B Me 7234). Vitis messelensis specimens are similar in overall morphology to the species V. hookeri Heer from the Oligocene of Bovey Tracey in southern England (Heer 1862, Reid & Reid 1910, Chandler 1957), corresponding well with the specimen illustrated by Reid & Reid (1910: pl. 15, fig. 3). Unfortunately, the only specimen of V. hookeri from Bovey Tracey, the type locality (NHMUK V33931 illustrated by Chandler 1957) is now very fragmented (c. 6 pieces) and it is not possible to observe any of the diagnostic characters other than a seed coat at least 150 µm thick and a prominent chalaza (not elongate but el-


liptical to circular). Chandler (1957) made a comparison between her specimen and those of Heer (1862) and Reid & Reid (1910) which makes it clear that V33931 was a newly collected specimen. Reid & Reid (1910) stated that their material was deposited in the Museum of Practical Geology, which has become the Geological Museum; the specimen is now housed in the British Geological Survey Keyworth, specimen number BGS CR1769. The specimen is very fragile and is glued onto a card such that it cannot be viewed from the ventral side (Howe, personal communication 2011) and the ventral side was not illustrated by Reid & Reid (1910). Therefore it is not possible to make a full comparison between any surviving specimens of V. hookeri and the Messel material. According to Heer (1862) the holotype measured 3.35 × 3 mm; but when more intact measured 3.75 × estimated 3.5 mm. Based on the stated magnification on the original illustration, the specimen figured by Reid & Reid (1910: pl. 15, fig 3) measured 3.75 × 2.8 mm but based on a new photograph provided by BGS the measurements of the specimen (BGS CR1769) are 4 mm × 3 mm. Therefore, it seems clear that V. hookeri is smaller than Messel specimens. Vitis glabra Chandler (1962) from the middle Eocene of Lake and Arne, UK, is also very similar to the Messel species is but it is smaller (maximum 4 × 2.75 mm) and has a slightly more prominent apical notch in the three known specimens. Overall, we have not found a close match among other fossil Vitis species and therefore describe the Messel material as a new species, V. messelensis. Specimens: Holotype SM.B Me 21619 (here designated, pl. 42a, b). Paratypes SM.B Me 145, 189, 456, 473, 2320, 2342, 2343–2346, 2347(inflated seed), 2370, 4025i, 4025ii (two seeds are in the same box and have the same number), 4151, 4209, 4216 (inflated seed), 4932, 4933 (group of 3 seeds), 4935 (inflated seed), 7121, 7122, 7123,7281, 7282, 7286, 7336, 7964, 7965, 7968, 7970, 7972, 7973, 7979–7982, 7984, 8210. SM.B Me 4746 is interpreted as a coprolite containing seeds of this species.

Genus Parthenocissus Planch. Parthenocissus britannica (Heer) Chandler (Pl. 42i–k, m–r) Basionym: 1862 Vitis britannica Heer, p. 1071, pl. 69, figs. 25, 26. Phil. Trans. Royal Soc. London, 152 (2): 1039–1086. Synonymy: 1957 Parthenocissus britannica (Heer) Chandler, p. 103, pl. 15, figs. 119–122. Bull. British Mus. (Nat Hist.) Geol., 3, 71-123, pls. 11-17.

Description: Seed obpyriform in dorsal and ventral view, (5) 6.0–7.2 mm long, (3.2)–4.0 mm wide, dorsiventral thickness ca. 2 mm (estimated from compressed specimens). Apex rounded without a groove or only faintly grooved. Beak pointed. Surface smooth but not shiny, not rugulate. Ventral infolds and dorsal chalaza of the endotesta somewhat obscured by draping exotesta. Ventral infolds linear, slit-like, straight, slightly diverging, relatively long, extending about two-thirds of the full seed length.

Chalaza central or offset somewhat apically on the dorsal surface, elongate-elliptical, occupying about one-quarter of the full seed length. Seed coat composed of a layer of anticlinally aligned cells 150–230 µm thick. Chalaza-apex and chalaza-base grooves weak. Fruit ovoid, 8 mm high, 7 mm wide, bearing two or more seeds. Comments: These seeds have long ventral infolds (pl. 42i, m) and an elongate elliptical chalaza (pl. 42j, k, n), but the details of these features of the endotesta are often concealed by the exotesta. The seeds are frequently compressed obliquely and it is not clear if they were originally symmetrical, or if this is simply a function of distortion by compression in the sediment. They resemble seeds of extant Parthenocissus in having long slit-like infolds that diverge apically, and an oval chalaza (Chen & Manchester 2007). Similar seeds from the Eocene London Clay and Clarno Formations have been assigned to the extant genus Parthenocissus (Scott 1954; Manchester 1994). For the London Clay species, Scott (1954) provided the new combination, Parthenocissus monasteriensis (Reid & Chandler) Scott. However, Chen (2009) found that these fossils, including the Messel material, differ from seeds of the extant genus in lacking the sharp apical groove (based on a survey sampling 8 of the 16 extant species). One of the species not sampled in that study, P. dalzielii Gagn., has seeds similar to the fossils, lacking the deep apical groove (our observations of B. Bartholomew 7749, from Taiwan). The traditional placement of fossil seeds within Parthenocissus needs to be revisited in light of the relatively recent recognition of Yua C. L. Li as a distinct genus. This extant Chinese genus, which is sister to Parthenocissus in morphological and molecular analyses (Lu et al. 2011), has seeds similar to Parthenocissus (illustrated by Lu et al. 2011 and Chen & Manchester 2011) but they lack the apical groove and are, in this respect, more similar to our fossils. Of the two extant species, Yua austro-orientalis (F. P. Metcalf) C. L. Li, is distinguished by pronounced rugulate ornamentation, while the type species, Yua thomsoni C. L. Li, has smooth-surfaced seeds similar in morphology to the Messel fossils. Lu et al. (2011) found that the ventral infolds tend to be shorter and more symmetrical and the seed base more acute in Yua than in Parthenocissus. It is possible that the Messel species represents Yua, rather than Parthenocissus, but a detailed investigation, including comparative anatomy of the extant and other fossil representatives was not completed at the time of this writing. These seeds coincide well in morphology with Parthenocissus britannica (Heer) Chandler, also known from the Oligocene and lower Miocene of NW Saxony (Mai & Walther 1991), but the Messel specimens reach larger sizes: mostly in the range of 6–7 mm long vs. 4.0–5.5 mm (Bitterfeld; Chandler 1957), and 4.75– 5.5 mm (Bovey Tracey; Heer 1862, Chandler 1957). Heer (1862) described the Bovey Tracey type material as 53


5.5 mm long and 3 mm wide and with indistinct chalaza – consistent with Messel material. The London Clay species, Parthenocissus monasteriensis (Reid & Chandler) Scott, also has a smaller seed, with internal cast 4.5 mm long and 3.5 mm wide (cited as Cayratia (?) monasteriensis Reid & Chandler 1933). Specimens: SM.B Me 2368 (loose from shale), 2369, 4102, 4213, 4220 (nicely broken), 4778, 4931, 4934, 5165 (broken showing seed coat anatomy) 5171, 5744 (fruit with two seeds exposed), 7337, 7974, 7976, 8262, 12271. SM.B Me 15840 Fruit with seeds.

Genus Ampelopsis Michx. Ampelopsis sp. (Pl. 43a–i) Description: Seed obpyriform in dorsal and ventral view, rounded apically, pointed basally but without prominent beak, 6 mm long, 4–4.5 mm wide. Ventral infolds 1/2 the length of the seed, diverging apically, with 6 or more ridges radiating distally from each of the infolds. Dorsal surface with 16–20 broad, sharp-crested ridges radiating from a centrally positioned raised elliptical or pyriform chalaza. Chalaza-apex groove absent or inconspicuous and chalaza-base groove weak, linear. Seed coat ca 170 µm thick, composed of cells in columnar arrangement. Comments: This species conforms to Ampelopsis in having widely divergent ventral infolds (pl. 43b, c, weak or absent apical groove, and spatulate chalaza (pl. 43a, d, g). Although many extant species of this genus, and the Eocene fossils, A. rotundata Reid & Chandler and A. rooseae Manchester (1994), have smooth seeds, this fossil is prominently ornamented with a pattern of radiating ridges similar to that of extant A. cantonensis (Hook. et Arn.) K. Koch (fig. 17 d in Chen & Manchester 2011). Chen & Manchester (2011) illustrated one of these specimens (SM.B Me 2352) as Ampelocissus cf. lobatum of Hordle, England, however the Messel material is distinguished by more widely divergent ventral infolds and barely protruding or not protruding beak. This species might correspond to Ampelopsis crenulata Reid & Chandler (1933) of the London Clay, but they are difficult to compare in detail because the illustrated London Clay seed is a locule cast lacking surface details of the seedcoat.

Synonymy: 1984 Ampelocissus jungii (Gregor) Gregor Documenta Naturae 16: 16, taf. 2, figs. 1, 2; 2007 Cayratia jungii (Gregor) Chen & Manchester, Amer. J. Bot. 94: 1548, fig. 11.

Description: Seed elliptical in dorsal and ventral view with subparallel convex lateral margins, 4.0 mm long, 2.8 mm wide, dorsiventral thickness unknown due to compression. Apex rounded, apical notch lacking. Basal side rounded, beak small, slightly protruding. Ventral surface fluted with grooves radiating from the raphe ridge. Ventral infolds wide, shallow and inconspicuous, symmetrical to each other, separated by a narrow median longitudinal raphe ridge. Dorsal surface with broad ridges radiating from the chalaza separated by shallow depressions. Chalaza elongate-elliptic, extending from the centre of the dorsal surface to the apex. Comments: This seed type is very rare at Messel. Broad ventral infolds are characteristic for some species of the genera Ampelocissus Planch., Ampelopsis and Cayratia. Of these modern genera, only Cayratia has an elongate chalaza extending fully to the apex as in this fossil (Chen & Manchester 2011). We are unable to distinguish this species morphologically from the younger specimens described as Cayratia jungii (Gregor) Chen & Manchester (2007) from the Middle Miocene of Schwandorf, Germany (Gregor 1977) and Oberdorf, Styria, Austria (Meller 1998). This genus has only rarely been recognised in the fossil record. Another occurrence of seeds of the same kind was called Ampelospermum pulchellum V.P. Nikitin (2006) from the Upper Oligocene to Lower Miocene of Dunayevsky Yar at Kompassky Bor on the Tym River, Tomsk region, Siberia. Specimens: SM.B Me 7613.

Genus Palaeovitis Reid & Chandler Palaeovitis sp. (Pl. 43j– n)

Genus Cayratia Juss. Cayratia jungii (Gregor) Chen & Manchester (Pl. 43o, p)

Description: Seed almost circular in outline in dorsal and ventral view, 4.3–6.5 mm high, 4.5–6.5 mm wide. Apex rounded sometimes with a very shallow raphe groove. Base rounded without a noticeable beak. Dorsal surface with spatulate chalaza, forming a ridge to the apex. Occupying about 1/4 to 1/5 of the width of the seed, with 12 to 13 high, rounded ridges radiating from the chalaza to the periphery. Chalaza-base and chalaza-apex grooves not obvious. Ventral surface obscured but showing a median ridge separating a pair of depressions interpreted as collapsed ventral infolds. Surface of testa with obvious fine cellular reticulate pattern formed of polygonal cell outlines. Broken ventral surface showing noncolumnar, apparently isodiametric, cells, composing the seed coat.

Basionym: 1977 Palaeocayratia jungii Gregor Paläontol. Zeitschr., 51: p. 214, abb. 8, 9, taf. 20, figs. 1–4.

Comments: This seed type, with its circular outline and radiating dorsal ridges, is similar in shape and size to

Specimens: SM.B Me 2352, 2339, 5166, 17395, 20381.

54


Palaeovitis paradoxa Reid & Chandler (1933) from the London Clay and Prémontré in Aisne, France (BlancLouvel 1986). None of the Messel specimens have been broken or sectioned in such a way as to clearly illustrate the seed coat anatomy, however oblique shattering of the ventral side of SM.B Me 7638, reveals isodiametric cells of the seed coat. Although Reid & Chandler (1933) indicated the chalaza of P. paradoxa is very large, this seems to have been based on a cast of the inside of the seed coat, rather than the external surface. Other specimens, both from London Clay and these from Messel, have a spatulate chalaza not out of proportion with those common among Vitis and other extant genera. The London Clay specimens are similarly rugulate on the dorsal surface (in some specimens, but smooth in others) and are similarly rounded in dorsiventral outline without a distinct beak and likewise lack an apical groove or have only a shallow one. One of the diagnostic features of this genus – the very thick seed coat documented in the London Clay specimens (Reid & Chandler 1933) – was not directly observed in the Messel specimens because they are intact, unbroken and not sectioned. However, when dislodged from the matrix, one of the specimens (SM.B Me 7638, pl. 43k) fractured obliquely through portions of the seedcoat, indicating at least moderate thickness, and composition of isodiametric rather than columnar cells (pl. 43n). In the London Clay specimens, cells of the testa are isodiametric, but arranged in columnar fashion (Reid & Chandler 1933, supplemented by our own observations).

Genus Crassivitisemen gen n.

Specimens: SM.B Me 4936, 7638, 10676.

Unnamed Vitaceae (Pl. 43q, r) Description: Seed obpyriform in dorsal and ventral view, 5.8 mm long and 4.8 mm wide, dorsiventral thickness unknown due to compression. Apex broadly rounded, with no notch. Base acute, beak not apparent, possibly broken. Ventral infolds narrow, slit-like, very elongate, diverging apically. Raphe ridge broad, with narrow median groove. Ventral surface striate; striae transverse across the raphe ridge and radiating apically and laterally outward from the ventral infolds. Comments: Comparable striated vitaceous seeds are absent from early or middle Eocene compression floras rich in Vitaceae seeds in England (e.g. Chandler 1962). It is possible that the striations are artifacts of animal gnawing (random non-paired grooves possibly resulting from insect gnawing) or perhaps of preparation of the specimen. Additional specimens are needed in order to confirm this ornamentation. Specimens: SM.B Me 4795 (dorsal view), 7983.

Type species: Crassivitisemen wildei (Chen & Manchester) comb. n.

Crassivitisemen wildei (Chen & Manchester) comb. n. (Pls 44a–m, 45a–d) Basionym: 2007 Ampelocissus wildei Chen & Manchester Amer. J. Bot. 94: 1547, fig. 8n–p.

Generic diagnosis and emended specific diagnosis: Fruit single-seeded, obovoid, bilaterally symmetrical, narrowly rounded basally, broadly rounded apically, but with a central apical depression giving a cordate appearance, 16–23 mm long, 11.5–13 mm wide, 6.5–8 mm thick. Fruit relatively irregularly rugulate to verrucate with a median longitudinal rib (vascular strand) on one face and a rounded basal scar. Pericarp 1.5 mm thick, composed of somewhat spongy tissue (but not much compacted in preservation due to rigid cell walls) of isodiametric cells. Epicarp a thin outer cuticle with isodiametric cells, hexagonal in surface view, 30–40 μm diameter. Epicarp surface slightly shiny and with slightly raised outer periclinal walls appearing like small papillae when well preserved. Mesocarp with thick outer layer of open thin-walled parenchymatous tissue and thinner inner layer of more compact tissue. Seed oval to pyriform in ventral or dorsal view, large, 11–14 mm long, 7–9 mm wide, dorsiventral thickness uncertain because of compression of the fossils. Apical notch and beak not prominent. Ventral surface smooth; ventral infolds linear, parallel to slightly divergent from each other, and about one-half to two-thirds of the full seed length. Dorsal surface rugose, with 7–8 pairs of ridges radiating out from the edge of the elongate oval chalaza. Chalaza located at the centre of the dorsal surface elongate, and about one-third of the full seed length. Chalaza-apex and base grooves present, linear. Seed coat very thick (1.2 mm), composed of nearly isodiametric cells aligned in anticlinal rows. Comments: This is the largest of the vitaceous seeds at Messel. The fruit SM.B Me 2297 has well preserved epicarp and there are multiple specimens of fruits with well preserved mesocarp indicating that the pericarp was more resistant to degradation than other Vitaceae, and demonstrating that they were consistently single-seeded. Chen & Manchester (2007) assigned this species to extant Ampelocissus based on surface morphology of the seed, without analysis of internal anatomy and without information on the fruit which bore these seeds. Now that the seeds have been recognised within fruits, and the internal anatomy of the seed has been investigated it is clear that the full set of characters does not support the assignment to this extant genus. Although the seed has the paired ventral infolds (pl. 45b, c) and dorsal chalaza (pl. 44j; 45a, d) diag55


nostic of Vitaceae, the seed coat (pl. 44g, i, l, m) is thicker than in any known seeds of extant genera of the Vitaceae. Palaeovitis from the London Clay flora and Messel have a similarly thick seed coat, but other features of morphology, like shorter ventral infolds, clearly distinguish Crassivitisemen from Palaeovitis. Most genera of Vitaceae have four or more seeds per fruit, unlike the single-seeded fruit of Crassivitisemen. Among extant genera of the Vitaceae, only Cissus L., and Clematicissus Planch. are regularly single-seeded (Chen & Manchester 2011). However both of these genera have an elongate chalaza that passes from the dorsal surface over the apical end of the seed, unlike this fossil which has an elongate-elliptical chalaza over the centre of the dorsal side of the seed. The composition of seed coat wall, made up of cells that are short (pl. 44m), rather than columnar, distinguishes this seed from those in the genera within the pentamerous flower clade (Ampelocissus, Vitis, Ampelopsis, Parthenocissus, and Yua C.L.Li), and suggests affinity with those with tetramerous flowers (e.g., Tetrastigma K.Schum., Cayratia, Cissus, Cyphostemma (Planch.) Alston) (Chen & Manchester 2011). Specimens: Holotype SM.B Me 7271(as designated by Chen & Manchester 2007); paratypes SM.B Me 7371, 7552, 8205, 8786. Additional specimens: Isolated seeds SM.B Me 2354, 4806, 4937, 5724, 5735, 5750, 7969, 8394, 17434, 17509, 17532, 18896, 21423, 21429, 21495, 21579. Fruits: SM.B Me 2297, 2298, 4648, 5727, 5729-5733, 5751 (fruit with ridges of the contained seed visible at one end), 8410 (nice cross section), 8418, 23885. SRXTM was applied to SM.B Me2298 which confirmed the seed morphology without destructive sectioning. Possible additional specimens: SM.B Me 4243, 4711, 5734, 23885 (these fruits have the same gross morphology but have not been dissected to confirm the seed morphology).

Incertae Sedis

Genus Carpolithus L. Carpolithus callosaeoides (Engelhardt) comb. n. (Pl. 45e–l) Description: Fruit ovate to widely ovate, massive, length 22–30 mm, width 18–25 mm, base rounded with a prominent bulging circular or almost circular scar 3–4 mm diameter, apex with a slight to marked conical depression. Surface finely longitudinally striate; striations radiating from the basal scar and converging apically. Fruit unilocular. Fruit wall thick (1.4–2 mm), composed of longitudinally elongate sclereids that are isodiametric in cross section 100–130 µm wide, with resin or latex secretions preserved in elongate strands. Resin fillings conforming to the outline of the cells. Locule usually collapsed due to compaction of the fruit, but containing a single seed (SM.B Me 2322). 56

Comments: We are uniting specimens that were called Ficus callosaeoides and Carya costata by Engelhardt (1922), together with more recently collected specimens. They are similar in size and share fine longitudinal surface striation, dimpled apex, thick sturdy wall, and prominent basal scar. The broken specimen (pl. 45k) shows a thick walled unilocular construction with a single seed, unlike Ficus. The fruit does not appear to have a plane of dehiscence or germination splitting, nor a prominent septum as would be expected in Carya and other Juglandaceae. The familial affinities remain uncertain, but the resin or latex material forming elongate strands as seen in the broken wall of SM.B Me 2322 (pl. 45l) may provide an important clue. Specimens: HLMD-Me-5333 [Carya costata Unger sensu Engelhardt 1922: pl. 32, fig. 12], HLMD-Me-2142 [Ficus callosaeoides type, Engelhardt 1922: pl. 12, fig. 6a, b], SM.B Me 2321, 2322, 4098, 14093.

Genus Saportaspermum Meyer & Manchester Saportaspermum kovacsiae Kvaček & Wilde (Pl. 46a–h) 2010 Saportaspermum kovacsiae KVAČEK & WILDE, Bull. Geosci., 85, 112-118.

Description: Seed winged, with an elliptical seed body, subrounded and strengthened at the base and somewhat pointed at the opposite end, with a single elongate membranous wing. The rounded base of the seed shows a relatively large circular scar. Surface of seed body showing a faint longitudinal striation from alignment of surficial cells. Seed body oriented obliquely to the long axis of the wing (30–40°). One of the lateral margins of the wing is straight, the other convex, and the distal margin is rounded. Wing membranous without obvious venation, except for a vein-like thickening on the straight margin. Wing not as prominently striate as the seed body. Most of the cells on surface of the wing are polygonal and normally isodiametric, but in occasional patches the cells are slightly elongate and aligned lengthwise to the wing, but striation not visible without magnification. Comments: There is some variability among this suite of specimens which may indicate more than one species. Similar seeds with a single elongate membranous wing occur in multiple angiosperm families, e.g. Malvaceae, Meliaceae, Proteaceae, Theaceae (Kvaček 2006). The fossil genus Saportaspermum is applied to seeds of this kind for which systematic affinities are unclear. The large circular scar at the rounded base of the seed (pl. 46f, h) is an additional character, seen also in the North American specimens, that may help in determining the systematic position of these seeds. This closely resembles Saportaspermum dieteri Walther & Kvaček from the Early Oligocene of Kundratice, Bohemia (Walther & Kvaček 2007). The Seifhennersdorf material is described


as having faint reticulate venation which we do not see in the Messel specimens, and the Messel specimens have one slightly thickened margin to the wing not described for the Seifhennersdorf material. Together with comparable material from from the Middle Eocene of Lábatlan, Hungary, and the Late Eocene of Kučlín, Bohemia, Czech Republic, the specimens from Messel have been assigned to a distinct morphospecies of Malvaceae by Kvaček & Wilde (2010), but their systematic affinities still need to be confirmed by more detailed comparisons. Specimens: SM.B Me 5576, 5587 holotype: Pl. 46e), 8586, 16455, 16836, 17019, 17020, 17447, 17836, 19885, 19950, 20181, 20616, 20349, 21620, 21622, 21809, 21810.

Genus Spirellea Knobloch & Mai Spirellea sp. (Pl. 46i–m) Description: Seed ellipsoidal, 4.0 mm long, 2.7 mm diameter, nearly circular in polar view. Slightly asymmetrical. Ornamentation consisting of 19 longitudinal sharp ridges meeting at both ends, ridges transversely striated, with closely spaced striae that are more pronounced between the ridges. There appears to be a smooth elliptical scar at one end and a conical less ornamented area at the other end. Faint longitudinal striations run longitudinally down from the elliptical scar and might possibly represent a raphe. Comments: This specimen is preserved uncrushed in three dimensional condition, not compressed, contrasting with many Messel fruits and seeds and indicating a rigid construction. It appears to fit with Spirellea a morphogenus to which Knobloch & Mai (1986) attributed 17 species from the Upper Cretaceous of central Europe. The shape, small size, longitudinal ribbing, striated ribs, numbers of ribs and terminal structures all agree. The Messel specimen, however, is larger than any of the numerous Cretaceous species. The Cretaceous seeds reach a maximum length of 2.4 mm, whereas the Messel specimen is 4 mm. The extension of this genus into the Tertiary has been indicated previously by Martinetto, who included it in a list of taxa (not illustrated) from the early Pliocene of Italy (Martinetto et al. 2007). The affinities of Spirellea remain to be demonstrated with certainty. Although Knobloch & Mai (1984, 1986) first treated this genus as Incertae Sedis they subsequently listed some of its species under the monocot family Stemonaceae (Knobloch & Mai 1991), but without explanation. In the course of comparative work with extant taxa, we observed disseminules of similar shape with similar ornamentation (straight meridional ribs and closely spaced transverse striae) in achenes of Fimbristylis annua Roem. & Schult. (Cyperaceae), Xyris pauciflora Willd. (Xyridaceae), and in the crassulaceous species Hylotelephium spectabile (Boreau) H. Ohba,

Kalanchoe spathulata DC., and Sedum latiovalifolium Y. N. Lee. Hence, there is convergence toward this kind of surface ornamentation in unrelated families. Reliable familial assignment will require more detailed comparative investigation. Specimen: SM.B Me 2196.

Carpolithus sp. 1 (Pl. 47a–f) Description: Fruit globose to obovoid with 10–12 straight, longitudinal grooves and a prominent basal attachment scar (about 2 mm in diameter) surrounded by a raised rim. Surface smooth at the macro level, but densely verrucate when viewed at higher magnification. The longest dimension ranges from 15–16 mm, and the shortest from 12–14 mm. Splitting from the apex along longitudinal planes into valve-like units. Wall about 1.2 mm thick, with outermost layer, 120–200 µm thick, composed of columnar cells, underlain by a less sclerotic layer 1.1 mm thick recognised by radiating fibers. Seed shape, size, and number not discernable, but tentatively a single seed revealed by patches of striate cuticle with striae parallel with the longitudinal axis of the fruit. Comments: We have been unable to locate any other modern or fossil taxon of similar size having such a large basal scar (pl. 47a, c, d) and prominent meridional grooves. Specimens: SM.B Me 7095, 7334, 14514, 15049, 17607, 18096, 19836, 20733, 21268, 23958.

Carpolithus sp. 2 (Pl. 47g–l) Description: Fruit with an elliptical seed body and a prominent long style. Style up to 17.5 mm in length, and 0.1–0.2 mm in diameter. Tissues continuous with style enveloping the basal body (0.2–0.5 mm thick). Basal body fusiform elliptical (length 5.0–5.2 mm, width 1.7–2.2 mm; n = 2), finely longitudinally striate with transverse ridges forming a faint reticulum. A small circular scar at the apex of the fusiform body may represent the micropyle. A small circular scar in the outer tissue at the base may represent an attachment scar. No indication that any hairs were present. Comments: A similar fossil was identified to Clematis vectensis (Reid & Chandler 1926) from the Bembridge Marls (Insect Limestone flora, Collinson et al. 2010) of England. These fossils correspond in size and shape and have a similar surface texture that Reid & Chandler (1926) described as having “large square cells aligned in 57


rows, and striate over the middle with small cells aligned longitudinally”. However, Reid & Chandler observed that “a cylindrical vascular strand enters at the base and immediately bifurcates, a branch passing up within the rim on each side; at the apex the branches re-united and enter the awn; at the apex of the locule one of the bundles gives off a short downward directed branch to the placenta of the seed”. These characters are not evident on the Messel specimens so it has not been possible to determine if the Insect Limestone and Messel specimens are the same species. Although these fruits resemble those of extant Clematis (Ranunculaceae) in shape of the ovary and long persistent style, extant species of Clematis are clothed in long hairs that occur both on the fruit body and the style, whereas these fossils appear to have been glabrous. Geum L. (Rosaceae) also produces long-styled fruits of similar shape, but they, too, are distinguished from this fossil by the presence of hairs on the fruit body and style. Specimen: SM.B Me 2211, 7605, 7712, 17776.

Carpolithus sp. 3 (Pl. 48a–i) Description: Fruit obovate, elongate, asymmetrical, sometimes D-shaped, with one lateral margin more or less straight to weakly convex and the other convex, length 53–76 mm, width 16–26 mm. Apex bluntly rounded, base truncate. Wall fibrous, sometimes giving a striate surface appearance, particularly evident near the truncate base, especially in the larger specimens. Surface smooth (pl. 48a, d) to coarsely transversely corrugated (pl. 48b). The external surface is densely covered with fine simple, black, unbranched hairs (pl. 48h, i). Comments: These seem to be trivalvate fruits, judging from an example that has been freed from the matrix, showing both sides (SM.B Me 2325, pl. 48d, e). Most specimens are very thin and appear to represent dispersed individual valves. Under high magnification, all of the specimens show a dense covering of fine hairs (pl. 48h, i) which completely cover the outer surface in the larger specimens (SM.B Me 7068, pl. 48b), but are preserved only in patches on the tripartite specimen (SM.B Me 8807, pl. 48h). One specimen (SM.B Me 8807, pl. 48f) shows three closely associated fruits, possibly attached, that are similar to the larger isolated fruits, and possesses the same kind of surface hairs. Its fruits are only 1/2 to 2/3 the length of the other specimens, suggesting that this may represent a cluster of immature fruits. These fruits resemble, to some extent, those of the Zingiberales on the basis of general shape and texture, but there are insufficient diagnostic characters available, in particular placentation type and seed anatomy. Specimens: SM.B Me 2325 7065, 7068, 7070, 8807 (small), 13738.

58

Carpolithus sp. 4 (Pl. 48j–n) Description: Seed ovoid to globose, 6.5–9 mm in diameter (n = 2). Relatively three-dimensional and shiny, with distinctive rectangular to polygonal cell surface pattern, formed from raised anticlinal cell walls, readily visible in reflected light microscopy; polygons vary from equiaxial (15–40 µm diameter) to elongate reaching up to c. 70 µm in length and giving a striate appearance in some parts of the seed (possibly overlying a raphe). Outermost seed coat layer very thin, cuticular (max. c. 40 µm), underlain by a thick layer (c. 350 µm) of parenchymatous spongy tissues. No scars or features seen on exposed part of seed. Comment: The seed coat surface of this species is clearly distinct from that of other shiny seeds from Messel. Carpolithus sp. 4 lacks the surface polygons enclosing small pits of Carpolithus sp. 57, and it has a well defined surface cell pattern unlike Carpolithus sp. 58. It is distinguished from Carpolithus sp. 39 by lacking surface hexagons interspersed with small open pits seen as depressions in LM. In addition, the inner seed coat layers of Carpolithus spp. 39, 57 and 58 include very thick-walled tissue (in some cases columnar) which differs from the more parenchymatous spongy tissues of Carpolithus sp. 4. The seeds are strongly crumpled and folded, suggesting ductile deformation unlike Carpolithus sp. 57. Specimen: SM.B Me 5057, 21562.

Carpolithus sp. 5 (Pl. 49a–c) Description: Fruit elliptical, possibly elongate, 28 mm wide, and at least 39 mm long, rounded at one end and broken irregularly transversely at the opposite end, and with rounded lateral margins. Fruit also broken longitudinally, revealing smooth locule(s). Fruit outer surface finely verrucate and with thin hairs sometimes in groups. Fruit wall thickness not known. An undulating longitudinal fibre bundle (2 mm wide in centre of fruit, broadening to 3 mm or more) extends down the centre of the specimen (possibly placental). Wrinkled shiny iridescent cuticle, with an isodiametric cell pattern, spreads across much of the exposed inner locule(s), on both sides of the central fibre bundle. In some places this cuticle appears continuous, in others it has straight to curved margins, and indications of overlapping layers, suggestive of overlapping membranous seed wings. Locule has widely ellipsoidal to rounded quadrangular shallow impressions (c. 2.5–3 mm in maximum length and 1.5–1.8 mm in breadth) that appear to represent original positions of seed bodies filling the locule(s). Seeds numerous, at least 26 impressions visible. Remnants of the seed coat are present in at least seven seeds (pl. 49b, c). Outermost seed coat layer secretory with pale infilling in


elongate or isodiametric cells, inner seed coat layer c. 90 µm in thickness, not columnar, black, shiny, with isodiametric cells c. 20 µm in diameter. Comments: The distinct notch in this fruit may be original but an artifact due to breakage during collection can not be ruled out. This fruit is superficially similar to the bignoniaceous pod valves that we referred to Darmstadtia, but is readily distinguished by the multiple rows of irregularly arranged seeds (rather than two distinct rows), the absence of a longitudinally striate margin of dehiscence, and much larger size. Although similar in shape to the fruits of Carpolithus sp. 3, this specimen is much larger than those and the internal features exposed in this broken specimen cannot be compared with the external surfaces known from Carpolithus sp. 3. We interpret this specimen as an immature capsule containing winged seeds. We only have this single incomplete specimen with no counterpart. Possible affinities include Meliaceae (if originally 5-partite) or monocotyledons (if originally 3-partite). The seeds (pl. 49c) appear to have had axile rather than parietal placentation indicating that this fruit does not belong in the Salicaceae/Flacourtiaceae but could be monocotyledonous (e.g. Zingiberalean), or malvalean.

(Schaarschmidt 1992: fig. 38); however, the discovery of specimens showing multiple seeds or pyrenes, rather than a single endocarp (pl. 49e, f), excludes assignment to that family. The inner elliptical bodies appear to represent seeds, although there is a possibility that they are pyrenes. The surrounding pericarp tissues were presumably very labile as only the cuticles have survived. The general organization of this fruit is similar to that of Canellaceae, but we have not undertaken detailed comparative analyses. Specimens: SM.B Me 4114, 4125, 13028, 14768.

Carpolithus sp. 7 (Pl. 49h–k) Description: Fruit a globose to oblate capsule, 6–7 mm high, 5.8–7 mm wide, with 3 bipartite valves delimited by longitudinal grooves. Surface finely verrucate. Pedicel stout, sometimes dispersed with the fruit, 0.8–1.1 mm wide, 3.5–4 mm long. Styles often persistent, at least 2 (perhaps a 3rd is fallen or hidden in sediment?), 1.2– 1.5 mm long, arising from a common point at the apex, widely recurved.


Carpolithus sp. 6 (Pl. 49d–g) Description: Fruit globose to oblate, 9–12 mm high, 9– 12.5 mm wide, borne on a prominent, longitudinally striate, non-woody pedicel 1.4 mm wide, 14–15 mm long; pedicel with central strand of thicker tissue enveloped by a cuticle (cuticle also forming a compression rim either side of pedicel). Pedicel cuticle with striations formed from fine narrow ridges (c. 10 µm) resulting in ‘stripes’ that extend the full length of the pedicel. Pedicel expanding both at the base and at the junction with fruit (to c. 1.8 mm). Attachment scar on fruit (seen due to displacement) oval, finely scabrate. Fruit strongly compressed (almost flat), covered by a crumpled tough orange to red cuticle (translucent in places); inner and outer cuticle surfaces show a pattern of polygonal isodiametric cell outlines (c. 30 µm diameter). Beneath the outer cuticle are 1–3 (or more?) flattened smooth elliptical bodies, 4 × 2 mm (seeds or pyrenes), with black, shiny cuticular surface. The outer surface of the black cuticle shows elongate cell outlines up to 80 µm in length and 20 µm wide. Comments: These specimens represent a berry-like fruit with a prominent well-preserved external cuticle. The enlargement of pedicel at the junction with the fruit suggests the position of a shed hypogynous perianth. Based on the distinctive resistant cuticle and drupaceous appearance, this fruit type was formerly suggested to be Lauraceae

Comments: Based on the morphological similarity but much smaller size, these capsular fruits might be immature or abortive examples of the species we have named Euphorbiotheca gothii. None of those fruits show the persistent styles which are typically preserved in these specimens (e.g. pl. 49h–k). Specimens: SM.B Me 8037, 8057, 8539, 14666, 18145.

Carpolithus sp. 8 (Pl. 50a–c) Description: Disseminules pyramidal with probably 4, or possibly 3, facets; ovate in face view, either quadrangular or triangular in cross section (symmetry uncertain because specimens still partially obscured by sediment at the time of this analysis); 2.1 mm long, 1.5 mm wide. Apex acuterounded, base truncate-rounded. Margins of adjacent facets joining to form smoothly rounded corner ridges. Facets beset with many small irregular dimples except at the smooth margins (pl. 50a, b). Surface formed by isodiametric cells arranged in rows giving a finely striate appearance, with striae longitudinally to obliquely arranged over the central portion of each facet, but more or less horizontally arranged over the corner ridges (pl. 50c). Comments: This fruit or seed type is known from three individuals, clumped together on the same piece of shale (pl. 50a). There is some similarity to fruits of Trema (Celtidaceae; Manchester 1989a), but detailed comparative work has not been done. 59



Carpolithus sp. 9 (Pl. 50d, e) Description: Disseminule widely elliptical in face view, 4.0 mm high, 4.2 mm wide with a softer, perhaps leathery layer enveloping a central endocarp or seed that is almost circular in face view, 3.0 mm long, 2.9 mm wide, bisymmetrical with a median ridge separating a pair of smooth, D-shaped concavities. Dorsal surface of the seed or endocarp obscured due to coverage by the fruit but suggestive of a smooth surface, a central depression and a median linear groove. Comments: The specimen has morphology and symmetry suggestive of being one unit of a bilocular capsule, or perhaps a seed of Vitaceae with wide ventral infolds surrounded by fruit tissue. The reverse side of the seed/ pyrene is obscured by persisting fruit tissue, making it difficult to obtain sufficient characters to identify the single specimen at this time. From the exposed features it seems to be distinct from all other Messel taxa we have examined, but it resembles a fruit type that is relatively common at Eckfeld (VW personal observations). Specimen: SM.B Me 16405.

Carpolithus sp. 10 (Pl. 50f) Description: Fruit a berry nearly circular in obliquetransverse view, 3.5 mm diameter, est. 3.0 mm high, containing 5 narrow cuneiform D-shaped pyrenes or seeds 2.4 mm high, 1.3 mm wide, and 0.3 mm thick, radiating from the central axis of symmetry. Fruit tissue collapsed, but surviving remnant indicates a smooth rounded surface. Comments: This obliquely compressed specimen shows clearly five radially arranged seeds (or pyrenes?). The rounded outline of the enclosing cuticle survives, indicating that other tissues of the fruit were readily degraded, probably soft or fleshy. Specimen: SM.B Me 13973.

Carpolithus sp. 11 (Pl. 50g) Description: Circular aggregation, 2 mm diameter, of approximately 13 wedge shaped units arranged around a deep central circular depression. Wedges 0.3–0.5 mm wide, variable in breadth, rounded externally, separated 60

from one-another by grooves. Surfaces radially striate with elongate cells up to 80 µm, long. Internal tissue of the wedge apparently consisting of isodiametric cells. Internal organization unknown. Comments: We cannot ascertain the depth of the grooves separating the wedges of this object, so it is possible that this specimen represents a single circular structure with surficial grooves, rather than a structure of distinct appressed wedges. Specimen: SM.B Me 5586.

Carpolithus sp. 12 (Pl. 51a–h) Description: Infructescence branching in at least two orders, with an articulation and small bracts at the base of the pedicel, internodes 4–5.5 mm in length. Pedicels 3–4 mm long. Fruit a wide-obovate capsule, 8.0–8.8 mm high, 8.0–10.5 mm wide (n = 3), widest in the upper third and narrowing to the pedicel, with convex sides, truncate apex and acute to right-angle base. Stylar protrusion preserved in one specimen 0.4 mm long (pl. 51b). Three visible free rounded hypogynous calyx parts 1.5 mm long (pl. 51b) (their arrangement indicates a total of five or perhaps six). Longitudinal ridges with central narrow grooves (pl. 51b) may indicate the edges of valves or carpel sutures. Thickness of the fruit wall as measured in a dehisced fruit (pl. 51c) 1.3 mm. A prominent axial strand (0.4 mm in width) extends from the pedicel into the locular area (pl. 51c). The strand apparently divides at the base (basal placentation) of the locular area with branches extending toward the seeds. One branch ends at a clear attachment to the seed (hilum); the other seems to pass underneath the seed in median plane of fruit. Seeds at least two per fruit, positioned in the apical three quarters of the fruit. Seeds elliptical in face view, inferred to be lenticular in cross section. In situ seeds 4.8–6.0 mm long, 3.0–3.1 mm, isolated seeds 5.4–6.3 mm long, 3.4–4.2 mm wide. Seeds with distinctive ornamentation of more or less longitudinally arranged distinct papillae (pl. 51d–h) which result from the upraised outer periclinal walls of the square surficial cells. No obvious micropylar, hilar or chalazal scars and raphe ridge have been observed. Comments: This species is represented mostly by isolated seeds easily recognised by their finely papillate surface ornamentation (pl. 51d, g, h). Two of the fruit specimens show these seeds in situ (pl. 51c, e, f) thus linking this seed type with fruit morphology. One of the specimens has dehisced one or more valves, indicating capsular dehiscence, clearly exposing the seeds, as well as the smooth marginal contact surface of the remaining valve, the central placental bundles, and two attached seeds (pl. 51c). The placement of longitudinal ribs (two


seen in pl. 51b) and configuration of the remaining valve of dehiscent fruit (pl. 51c) suggests a tricarpellate fruit. Useful characters for the systematic assignment of this taxon include the hypogynous calyx with well defined lobes (estimated to be 5 or 6 lobes, pl. 51b), the central, apparently single, persistent style (pl. 51b), the likely tricarpellate capsule with few seeds, and upraised cells of the seed coat giving the papillate ornamentation. Affinities might lie with the Ericales near Theaceae, but we have not determined its precise relationships. Specimens: Fruits SM.B Me 7096, 7235, 10447. Seeds SM.B Me 284, 4171, 4211, 4451, 4463, 4504, 4636, 4790, 7588, 7591, 7592, 8541, 8702, 8716, 8732, 14070, 20089, 20292.

Carpolithus sp. 13 (Pl. 51i, j) Description: Irregularly globose, valvate fruit, 33 mm long, 33 mm wide, representing a leathery longitudinally dehiscent capsule. Outer surface irregularly verrucate. Locules relatively smooth. A central axis of multiple strands extends the full length of the fruit, convexly deflected in the central third of the fruit length. Prominent disk, without obvious scars, at base of fruit, 6 mm long and 11 mm wide having ring-like structure at its base. Pedicel stout (c. 6–7 mm thick). Seeds not preserved. Comments: The number of valves within this fruit is uncertain; two are remaining, but the number of shed valves on this specimen could have been two or three. We thus may infer either a 4– or 5-carpellate fruit. The facing valves of the fruit have been shed exposing the internal locular surface. The deflexed central strands indicate possible site of placentation. If the disk represents the site of attachment for perianth parts, this would have been derived from a hypogynous flower. Specimen: SM.B Me 7064.

Carpolithus sp. 14 (Pl. 52a–f) Description: Irregular cluster of more than 46 obovoid to pyriform seeds, broadly rounded at one end and obtusely pointed at the opposite. Seeds 1.4–1.8 mm long, 0.8–1.4 mm wide. Seed surface irregularly corrugate (pl. 52c, b), surface composed of isodiametric pentagonal to hexagonal cells circa 7–8 µm in diameter. In places the outer periclinal wall is preserved, in others it has been broken or abraded to reveal thick walled cells with subangular lumen c. half of the cell diameter. Abrasion also reveals tiny cavities within the cell wall (pl. 52f). Remnants of cuticle are preserved along the anticlinal walls of the epidermal cells (pl. 52e).

Comments: This cluster of seeds could be interpreted as the remains of a multiseeded fleshy fruit, or as a coprolite. We prefer the former because it contains only one seed type, has indication of a radial arrangement of the seeds suggesting original placentation, and has continuity of a partially surrounding organic layer. Fleshy fruits with abundant elliptical seeds occur in multiple families. Moraceous fruits, for example are distinguished by seeds with smooth surfaces and typically an indurated funicular process (Collinson 1989b). Sambucus seeds are widespread in the Paleogene and are known from sieved material from Messel site 7. The size agrees with that of other Paleogene Sambucus, e.g. S. parvula Chandler 1926 (Collinson 1983b). However, the surface anatomy is quite distinct from Sambucus. The Paleogene Sambucus seeds have a more pronounced horizontal corrugation, and epidermal cells that are twice as big and have up to eight-sided cells, with a small papilla at each angle. Sambucispermites Manchester (1994) is similar in size and shape, but differs by having strongly horizontal corrugations. Specimen: SM.B Me 4086.

Carpolithus sp. 15 (Pl. 53a–d) Description: Fruit a large spherical capsule, 44 mm in diameter, with four distinct equal closed valves delimited by meridional grooves extending from near the base fully to the apex. Surface ornamented with evenly distributed, well spaced, short, broad-based blunt spines (2.0–2.5 mm in length, 1.0–1.4 mm in diameter). Comments: This species is known only from a single well-preserved specimen, but is very distinctive. Large spherical ornamented valvate fruits are known in extant Flacourtiaceae, e.g. Lindackeria. Other families with large globose spiny valvate fruits include e.g. Sapindaceae (some Aesculus), and Elaeocarpaceae (e.g. Sloanea). This is unlikely to be Aesculus which typically has three valves. Extant Sloanea fruits vary from 3 to 5 valves and often have more fragile and narrower, more closely spaced spines. Because of the orientation within the sediment we were unable to observe the base of the structure. Thus, it remains unknown whether there was a prominent basal disk as in Aesculus and Sloanea. Specimen: HLMD-Me-13399.

Carpolithus sp. 16 (Pl. 53e–g) Description: Branched infructescence bearing elliptical, longitudinally striate fruits, 9.0–10.3 mm long, 61


3.9–4.3 mm wide. Fruit with a hypogynous encircling rim interpreted as perianth scar; without obvious style. Fruits longitudinally fibrous. Comments: This specimen was initially grouped with the infructescences of Carpolithus sp. 29, because of similarities of overall branching pattern and fruits having hypogynous perianth scar. However those fruits are somewhat larger, have more broadly rounded apices and have a smooth texture unlike the striate texture seen in this specimen. Specimen: SM.B Me 11013.

Carpolithus sp. 17 (Pl. 54a–d) Description: Branched infructescence with numerous ovoid to globose individual fruits, and bracts subtending the nodes of branching. Pedicels articulated 6–7 mm below each fruit, prominently flaring and conical beyond the articulation to join the disk-like structure at the base of each fruit. Fruits 6.0–6.5 mm long (measured from top of disk to fruit apex), 4.5–5 mm in equatorial diameter, rounded basally and apically, with a circular bulge 0.3 mm high and 1.2–1.9 mm in diameter at the apex. A prominent disk-like structure at base, 3.5 mm wide, divided into regular, rectangular facets – 3 of these in face view (estimated 5–6 total) representing an abscised perianth and/or stamens. Comments: The distinctive characters of this taxon are a conical inflation of the pedicel, a conspicuous hypogynous disk, and an apical circular bulge (pl. 54b, c). This specimen was figured previously as Theaceae (Schaarschmidt 1992) but this was based on the cuticle of the leaves on the same slab, which cannot be demonstrated to be attached but appear to be simply overlapping. The unusual conical thickening of pedicels is reminiscent of the hypocarps in some Anacardiaceae. These fossils are clearly distinct from the curved fruits of Anacardium, but similar to Semicarpus fruits (see Manchester et al. 2007: fig. 2a–c). Some of the characters of this taxon have also been found among tropical tilioid taxa (Malvaceae) (Dilcher & Wilde, unpublished), but further comparative work is needed for a well-substantiated identification. Specimen: SM.B Me 2733.

Carpolithus sp. 18 (Pl. 55a, b) Description: Fruit large, 64 mm long, 46 mm wide, outline elongate-cordate with a distinct sinus at one end, split 62

longitudinally during collection to reveal seeds but with no evidence of a natural plane of dehiscence. External surface exposed near base essentially smooth. Probably with a single large locule; number of carpels unknown. Fruit wall about 4 mm thick. Fruit texture suggests originally leathery to fleshy rather than woody. Seeds at least 3, medially positioned in the fruit. Seeds large, collapsed, 16 mm high, 19 mm wide, slightly shiny, black, fracturing and readily detaching, with thin outer cuticular layer, and irregular longitudinally striate appearance due to arrangement of quadrangular to polygonal epidermal cells. Underlying the cuticular layer is a substantial spongy textured cell layer within which are longitudinal fibrous strands. Possible hilar scar is present as a discrete patch of spongy tissue surrounded by a well marked elliptical patch from which cuticular layer radiates. Comments: This is among the larger fruits from Messel. Distinctive features include the large somewhat shiny cuticle-covered seeds, and sinus at one end of fruit. But, it can not be completely excluded that the sinus is caused by accidental breakage. Affinities remain unknown. Specimen: HLMD-Me-16934.

Carpolithus sp. 19 (Pl. 55c, d) Description: Fruit pedicellate; pedicel 3.8 mm long and 1 mm thick, with a disk-like structure, 3 mm wide, and 1 mm thick, at junction with base of the fruit. Fruit widely obovate, length excluding basal disk 9.5 mm, width 12.3 mm, apparently membranous but without obvious venation. Margins of the fruit smooth and convex, no obvious stylar or stigmatic protrusion. In face view, a lateral arched linear structure appears on the fruit surface, possibly representing the edge of a wing. A single smooth obovate seed, 5 mm long and 5 mm wide, relatively thin walled, is preserved within the margins the fruit, possibly in situ. Hilum and placentation unclear. Comments: The mode of dehiscence and the stylar conditions of the fruit are unknown. It is uncertain if the seed is in situ, and therefore belonging to the fruit, or may have been deposited separately on the fruit and representing another taxon. The seed is similar in shape and size to those of the Messel Magnolia species, but appears to be more symmetrical and is lacking expression of the terminal cleft. The general organization of this fruit, smooth with hypogynous expanded perianth and/or disk scar, possibly membranous texture, and smooth elliptical seed (if the seed indeed belongs to this fruit), suggests affinities with Brassicaceae, Sapindaceae, Rutaceae, or Zygophyllaceae. Specimen: SM.B Me 17403.


Carpolithus sp. 20 (Pl. 55e, f) Description: Fruit widely elliptical outline, but with a retuse apex; length 16 mm, width 15.2 mm; smoothly convex laterally with remnants of at least two protrusions in the apical cleft. Pedicel 3 mm long and 1 mm thick, basal hypogynous disk or perianth scar 2.5 mm wide and 1.2 mm long. At least four wings may be seen, radiating from the median axis of the fruit, each with a strong marginal vein fed by subparallel veins that radiate from the midline of the fruit. Judging from preservation, the wings probably were chartaceous. Seeds not observed. Comments: The veins of the wings on this specimen are not immediately obvious because of the uniformly dark preservation. However, the free part of the outer right wing has a network of veins joining orthogonally to the marginal vein (pl. 55f). The fin-winged structure of this occurs in numerous extant angiosperm families, but the combination of four or more wings with a very stout pedicel, hypogynous perianth scar, strong marginal vein, and relatively widely spaced main veins spreading across the wings seems to be restricted to Malvaceae (Manchester & O’Leary 2010), e.g. to Craigia, Burretiodendron. Specimen: SM.B Me 8404.

Carpolithus sp. 21 (Pl. 55g, h) Description: Tiny winged seed. Seed body elliptical, 0.9–1.6 mm long and 0.45–0.7 mm wide, rugulate with transverse and longitudinal ridges probably caused by compaction and surrounded by a smooth membranous wing in the plane of compaction. Wing 1.2–1.8 mm long and 0.6–1.0 mm wide without visible veins. Comment: This taxon is one of the smallest disseminules in the Messel collections, possibly only recognised by collectors because of more than one specimen in close proximity. The same type of winged seed with cuticularized seed body, although still unnamed, is also known in collections from the Early to Middle Eocene of North America (e.g. Barrel Springs, Wyoming (UF1903053748), and White Cliffs of the Clarno Formation (UF 263-16929). These are at least superficially similar to the winged achenes of Machaeria (Cyperaceae; fig. 3 in Strong 1997). Specimen: SM.B Me 8708 (with two seeds of same type on the same block), 10508 (several seeds together).

Carpolithus sp. 22 (Pl. 56a–k)

Description: Fruit unilocular, ellipsoidal; height 14– 17 mm, width 10 mm. Ornamented with numerous stout spines arranged in longitudinal rows. Distinct rows of longer spines arranged on 4 meridional ridges. Shorter spines arising between the ridges, and on the lower flanks of the ridges. Most of the spines straight, unbranched, but some are distally bifurcate (as seen in SM.B Me 2059, pl. 56b, j). Longer spines narrowly conical, c. 0.7–0.8 mm, wide at base and sharply pointed, up to 2.6 mm long; shorter spines blunt-tipped. One of the specimens (HLMD-ME-16935, pl. 56d), with some of the spines represented only by their bases, has the spines arranged in distinct longitudinal rows separated by grooves (the median row is biseriate) [supported by what is seen in SRXTM of SM.B Me 2059]. Broken end of HLMD-Me-16935 exposes an underlying smooth surface with a median groove. Wall apparently uniform, 360 µm thick in areas between spines; formed of cells that are noncolumnar; spines show the same x-ray attenuation as the wall. Total fruit thickness 3.9 mm spine tip to spine tip, measured perpendicular to the plane of compression in SRXTM of SM.B Me2059. Comments: These are the only specimens with such robust spines known from the Messel collections. Specimens are typically exposed in the shale with only one surface visible so that the distribution of spines over the complete surface was uncertain until one of them (SM.B Me 2059, pl. 56b) was studied by SRXTM. The resulting digital sections (pl. 56e–k) reveal features that would not have been visible without destructive fracturing and/or physical removal from the shale. The SRXTM observations confirm that the spines protrude in all directions, that there are four major spine-bearing ridges, and that there is a single longitudinally elongate locule. Strong compaction makes it difficult to determine whether the locule was originally circular vs lenticular in cross section, and whether the overall symmetry was bilateral or 4-fold. Among extant fruits commonly possessing such spines, Ceratophyllum (Ceratophyllaceae), may be eliminated by its smaller fruits with a more pronounced apical spine; members of Tinosporeae (Menispermaceae) have boat or cup-shaped endocarps with a large ventral hollow which we have not observed in these specimens. Several genera of Apiaceae have spiny fruits that may come into consideration, but the epigynous perianth that typically persists in those fruits is not seen in this fossil, and the asymmetry that would be expected in mericarps of Apiaceae is not present in this taxon. Fruits of Apiaceae are typically borne in pairs with a resulting asymmetry of each dispersal unit; whereas Carpolithus sp. 22 fruits appear to have been fully symmetrical. Specimens: SM.B Me 484, 2059, HLMD-Me-13925, 16935.

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Carpolithus sp. 23 (Pl. 56l, m) Description: Oblate capsule, obliquely compressed, estimated 4.0 mm high, 4.6–5.2 mm wide. Surface smooth at the macroscopic level, with a prominent recessed area 0.8–0.9 mm diameter at one end. Fruits composed of 3 radially arranged bipartite units which are equal in size and shape. Units 2.2–2.5 mm wide, splitting from one end (apical in SM.B Me 10588). Wall thin, c. 70 µm, composed of apparently isodiametric cells (not columnar). Internal structure of fruit and seeds unknown. Comments: The fruits may be inferred to be both loculicidally and septicidally dehiscent. The 3-fold symmetry suggests affinities to monocots or Euphorbiaceae. The fruit is distinguished from Euphorbiotheca gothii by its smoother surface, considerably smaller size and much thinner wall. Specimens: SM.B Me 4174, 10588, 20196.

Carpolithus sp. 24 (Pl. 56n, o) Description: Seed elliptical, 5.8 × 4.1 mm, with a median longitudinal groove and multiple (greater than 10) low irregular transverse ridges. Surface with dark, shiny cuticle-testa layer adhering in patches; the shiny layer seems to follow into the transverse grooves. Surface beneath the cuticle formed by isodiametric polygonal cells 20–25 µm in diameter, underlying tissues formed of cells about 10–15 µm in diameter. Comments: The surface of this seed shows a pattern indicative of underlying ruminate endosperm arranged in a bisymmetrical pattern common to seeds of Annonaceae. The seed coat was relatively thin, such that the surface formed transverse ridges is reflective of the underlying blades of seed coat penetrating the seed cavity. Superficially similar seeds are found, for example, in the annonaceous genera Klarobelia Chatrou and Oxandra A. Rich. (figured in Cornejo & Janovec 2010). However, we have not made the detailed comparative investigations necessary to verify the affinities.

Comments: This fruit is similar in size to Carpolithus sp. 23 but distinguished by shape which is longer than wide, and ornamentation which includes longitudinal ribs, rather than grooves. Specimen: SM.B Me 7352.

Carpolithus sp. 26 (Pl. 56q–s) Description: Seed elliptical, length 1.3 mm, width 1 mm, relatively smooth, surrounded by radiating extremely fine hairs, up to at least 2 mm length, that appear to arise from just one end of the seed. Although the hairs overlap as preserved, there is no evidence for branching or grouping in tufts. Comments: Initially we considered that these tiny hairy seeds might represent Salicaceae, e.g. Salix L. or Populus L., however seeds of the extant species of these genera are more narrowed apically, whereas these fossils are well rounded both apically and basally. Apocynaceae are also known for genera with tufted seeds, but the seeds of those genera are elongate, generally three or more times longer than wide. Specimens: SM.B Me 20154, 23128.

Carpolithus sp. 27 (Pl. 57a–d) Description: Raceme bearing at least 30 subsessile fruits. Infructescence 73 mm long, 16 mm wide. Fruits apparently spirally attached to a thick axis, 1.6–2.0 mm wide, fruits borne on peglike, very short broad pedicels, 0.8 mm long, 1.2 mm wide, which are left behind at shedding. Individual fruits globose to obovate with a fused stout cup-like basal half having straight vertical ribs (up to five in face view, estimated about ten in complete perimeter). Cuplike portion terminating in a transverse rim, giving rise to a whorl of five rigid tepal-like lobes. Lobes appear valvate, free from each other, and form a single whorl. The individual lobes are rounded triangular and have a network of pinnate veins arising from the transverse ridge. The fruits have no indication of a style.


Carpolithus sp. 25 (Pl. 56p) Description: Fruit ovoid, 5.6 mm long, 5.1 mm wide. Fruit with an irregularly rugulate to verrucate surface and six well-defined longitudinal vein-like ribs. Wall thickness and internal structure unknown. 64

Comments: The cup-like structure forming the basal half of these fruits appears to represent a hypanthium, from which persistent tepals arise. Normally two to three appear with others hidden from view, but a total of 5 can be counted in some. The architecture of the raceme and hypanthea resembles some Hamamelidaceae, e.g. Fothergilla, but the flattened, tepaloid structures do not resemble the perianth of Hamamelidaceae, and appear too thin to represent capsule valves. Possible affinities


with Rosaceae and or Hydrangeaceae have not been fully explored. Specimen: SM.B Me 7040.

Carpolithus sp. 28 (Pl. 57e–g) Description: Disseminule broadly elliptical, length 8.3 mm, width 7 mm mostly smooth, to slightly granular, without ribbing, but with a prominent convex basal scar marked by change in surface texture occurring abruptly at a transverse line at the lower 1/10 of length. Wall 0.2–0.4 mm thick. Comments: This species, known from a single specimen, shows differences in apparent surface texture, depending on whether it is observed in water (pl. 57e) or in dry condition (pl. 57f). SRXTM digital sections reveal that the disseminule has been flattened by compaction, that the wall is 0.2– 0.4 mm thick (thickest in the plane of compaction; estimated original thickness c. 0.3 mm), and now has numerous anticlinal cracks, probably resulting from shrinkage but with a single internal cavity (locule if a fruit, or embryo/endosperm cavity if a seed). Wall anatomy not resolved. A distinctive feature of this disseminule is the abrupt distinction in surface pigmentation and texture over the basal tenth of the length, resembling a large scar (hilar, aril, or perhaps cupular). Superficially, the specimen resembles a seed of Taxus L., with the basal truncation corresponding to the position of aril attachment, however, the fossil does not show the expected pair of vascular bundles (cf. Manchester 1994: pl. 2). There is a close resemblance to nuts of Coryloideae (Betulaceae), especially those of Carpinus L. and Palaeocarpinus Crane (cf. Manchester et al. 2004: fig 2), with the basal truncation representing the involucre scar, but we have not confirmed whether the wall anatomy corresponds. Specimen: SM.B Me 21351.

Carpolithus sp. 29 (Pl. 58a–g) Description: Infructescences (maximum preserved length 62 mm in SM.B Me 7540) with at least three orders of branching. Pedicels up to 3 mm long, abruptly widened at the junction with fruits. Individual fruits oblong-elliptical to obovate, length 6.6–9.0 mm, width 4.1–4.8 mm, rounded apically, and basally, with a prominent disk-like structure at base. Fruits relatively smooth, with 1–2 longitudinal ridges visible in face view (three in total), delimiting valves, that in some specimens are opened apically (pl. 58f). Individual valves have transverse wrinkles in the distal part and often bear a faint median groove.

The base of the individual fruits shows a stout thickening similar in shape and dimensions to the disk at the top of the pedicel, giving a “double” appearence to the basal thickening (pl. 58g). Seeds and placentation unknown. Comments: These fruits appear to be tricarpellate, judging from the symmetry of longitudinal ridges (pl. 58g), and a specimen clearly showing a dehiscent fruit with the three apically spread valves (pl. 58f). Possible affinities could lie with monocots such as Liliaceae, but dicotyledonous families, e.g. Salicaceae s.l. also share a similar kind of inflorescence branching and capsular fruits. Specimens: SM.B Me 2057, 2058, 7039, 7048, 7183, 7190, 7218, 7540, 8948, 14607, 14749, 15411, 15461, 15537, 16560, 16627, 17050, 17067, 17210.

Carpolithus sp. 30 (Pl. 58h–i) Description: Fruit ellipsoidal, length 33 mm, width 19.1 mm (l/w 1.9), with a distinct rounded apical protuberance 3.3 mm wide and 2 mm high. Fruit surface irregularly rugulate. Seeds (exposed by damage to a portion of the fruit) numerous, subglobose, 1.8–3.8 mm in longest dimension, with a circular depression at one end with a small protrusion of less than 0.5 mm diameter in the centre. One end of one seed shows a patch of distinctive tissue which appears papillate. The outer seed surface has a thin cuticle with rows of small rectangular cells, underlain by a thin black shiny layer; in turn underlain by two additional layers, both of which appear in reflected light microscopy to be composed of isodiametric cells. Comments: The seeds of this fruit are similar in their barrel shape and circular terminal depression to the operculate seeds of extant and fossil Musaceae. The circular depression resembles the area surrounding the operculum in modern and fossil Ensete Bruce ex Horan. (Manchester & Kress 1993). The seed apex looks similar to Carpolithus species 50 or 58 but is much smaller. The fruit appears to have been berry like, with a leathery pericarp rather than capsular. Although zingiberalean affinities appear to be likely, details of placentation and seed anatomy that would confirm this remain unknown. Specimen: SM.B Me 20155.

Carpolithus sp. 31 (Pl. 59a–k) Description: Seed ovate to subovate in face view, length 5.0–7.0 mm, width 3.3–4.3 mm (n = 6), the narrow end slightly bulging and thickened with an asymmetrical deep cleft (0.5–0.9 mm in depth). One face of the seed has a pair of broad longitudinal facets, 65


separated by a narrow ridge with a vertical vascular strand (raphe?) that runs from the base of the cleft to the broader end of the seed where there is a small indistinct scar. The opposite seed face is rounded and lacks vascular strands. Seed surface smooth, showing polygonal, more or less equiaxial, cells c. 50 µm in maximum diameter (pl. 59i) visible as dark shiny patches separated by lighter walls. Seed coat 90–100 µm thick; main thickness of seed coat formed by a layer of columnar cells (pl. 59j, k). Comments: The narrowed apex with prominent cleft is interpreted as the hilar scar with a raphe running along the median ridge towards a basal chalaza (pl. 59c, d, g, h). Specimen SM.B Me 4029 (pl. 59 c–e) was removed from the matrix, proving that only one side has the vascular strand interpreted as the raphe. Cells seen on the surface as polygonal, and more or less isodiametric (pl. 59i), appear to represent the proximal sides of cells that are columnar as seen in SEM of broken seeds (pl. 59 j, k). The seed morphology is consistent in many details with seeds of some Rhamnaceae, e.g. Rhamnus L. and Frangula Mill. Seeds in this family commonly have a seed coat of columnar cells, may have a prominent hilum, and seeds that are more or less trigonal, with a rounded dorsal face and more flattened ventral faces that adjoin a median ridge traversed by a raphe. In Rhamnus, the raphe runs medially down the dorsal surface, as in this fossil (Vyshenskaya 2000). In the seeds that we have examined (Hovenia Thunb., Rhamnus, Colubrina Frich-Joset & Montandon), the columnar cells of the seed coat appear to be much narrower than those of this fossil. Specimens: SM.B Me 2166, 4016, 4029, 4030, 5148, 5150, 8484, 10509, 17234, 17294, 18147, 19218, 19509. There is also a specimen on the block 7286 which also has a Vitis messelensis seed.

Carpolithus sp. 32 (Pl. 59l) Description: Disseminule ovoid (if pedicellate) or obovoid (if persistent style), 10 mm long, 7.1 mm diameter, narrowed and truncate at one end, rounded, with a stout protuberance at the other end, 0.5 mm thick and 2.1 mm long. No evidence of perianth scar at either end of structure. Longitudinal ridges extend the full length of the fruit. Ridges diminish in size towards narrow end. Eight ridges on the exposed lateral face, indicating a total of about 16. Comments: This specimen shows some resemblance to Doliostrobus cone scales (pl. 1a–c), but is smaller than typical specimens of that genus and it lacks resin. Specimen: SM.B Me 7079.

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Carpolithus sp. 33 (Pl. 59m–q) Description: Immature fruiting head, 9.5 mm long and 6 mm wide, with pedicel (2 mm length preserved, 0.75 mm in width) that expands slightly at attachment to fruiting head. A small protrusion, low and central on the pedicel, is a former scar of attachment of a lateral organ and a small transverse ridge higher and to the left side of the pedicel may indicate attachment of another (both possibly bracts). Base of head with possible overlapping bracts but individual components cannot be clearly distinguished and these may be compressed, aborted, non-developing fruits. About 68, probably spirally arranged, developing fruits visible on the exposed surface, suggesting up to 136 in the entire fruiting head. Developing fruits inflated at their bases, but variously irregularly indented and wrinkled, probably due to compression or shrinkage of poorly lignified tissues. In inferred transverse section the fruits would be oval to circular, or diamond-shaped where closely packed. Developing fruits towards apex of fruiting head narrowing to elongate attenuate apices. Maximum exposed attenuation length c. 4 mm narrowing gradually to pointed tips less than 100 µm wide. Attenuations clearly ductile near their tips as shown by bends and sinuosity. Some attenuations have a single very narrow longitudinal groove. No bracts are visible between the fruits. No fluorescing material (pollen or cuticle) is present on the specimen. Comments: The only comparable mature fruiting heads from the Messel oil shale that are in any way similar to this specimen are those of Volkeria (Cyperaceae) (pl. 8f–i; Smith et al. 2009b). Volkeria is estimated to have had between 16 and 70 floral units per spike with up to 35 visible on exposed portions, only about half as many as in the specimen described above. No specimens of Volkeria show any evidence of attenuations at the fruit tips and it seems unlikely that all evidence of attenuations in developing fruits would be lost in mature fruits. The lower developing fruits in SM.B Me 12158 do not show long attenuations. Some pointing outwards are clearly broken while those pointing sideways may have been damaged during preparation. It is possible, however, that the lower and upper units represent different structures possibly derived from different types of flowers with the upper units being sterile. Among the extant mapanioid Cyperaceae considered in comparison with Volkeria (Smith et al. 2009b) Lepironia Rich. fruits have apical attenuations (Smith et al. 2009b: fig. 6G) so an affinity with Cyperaceae is plausible for this taxon, too. An alternative possible affinity is with Araceae, especially if the lower and upper units are derived from different types of flowers. Araceae with similar infructescence morphology have fruits that are fleshy berries (see discussion in Smith et al. 2009b: 1515). In SM.B Me 12158 it is not possible to determine the fruit tissue structure or wall layers. However, the


wrinkling and indentations on the bases of the lower developing fruits might indicate shrinkage or compression of fleshy tissue. In the absence of other specimens and without mature specimens the systematic affinity of this specimen remains unresolved. MicroCT scanning was also applied to this specimen (pl. 59n–q). Although internal morphology is not resolved, these images show that the portion of infructescence still buried in shale conforms to the structure exposed at the surface, and that the structure was relatively resilient, showing the effects of compaction, but not completely flattened. Specimen: SM.B Me 12158.

Carpolithus sp. 34 (Pl. 60a–l) Description: Seed subelliptical in outline, slightly pointed at both ends. Length 7.3–9.2 mm, width 4.6–5.6 mm (n = 5). Nearly lenticular in cross section but with beveled edges. Surface is relatively smooth but with gently raised ridges forming a regular reticulum more radiating in the outer part, delimiting three- to six-sided areoles about 0.6–0.8 mm across. Wall about 160 µm thick. Surface covered in patches by polygonal isodiametric cells (pl. 60j); where that layer is removed, an underlying layer of thick-walled cells with sunken quadrangular lumen is prominent which gives a pattern of fine pitting when viewed by reflected light microscopy. Cellular construction of wall unclear due to diagenesis, but not markedly columnar. Comments: In one of the specimens (SM.B Me 7092, pl. 60e), three of the seeds are enclosed in a weak circular (originally globose?) structure with a faintly delimited margin. This represents either a coprolite or a fleshy fruit, but we favour the second interpretation because of the uniformity and continuity of the organic layer overlying the two exposed disseminules seen in pl. 60e, f. A similar reticulum and flattening in the plane of symmetry also occurs in endocarps of Icacinaceae (e.g. cf. Palaeohosiea, pl. 15). In Icacinaceae, however, the fruits are drupes in contrast to SM.B Me 7092, where there appear to have been multiple seeds per fruit. Also, the fruits of Icacinaceae would be more asymmetrical at the apex due to the presence of a funicle. Scanning electron microscopy shows the construction of seed coat, and impressions where rhomboidal crystals may have been borne (pl. 60g–l). Specimens: SM.B Me 2089, 4782, 7092, 7100, 16388, 24614.

Carpolithus sp. 35 (Pl. 60m–o)

Description: Cluster of at least four sessile, apparently ellipsoidal seeds or endocarps, 14 mm long, 9 mm wide, each covered by a thin layer with longitudinally oriented venation occasionally anastomosing, forming elongate, coarse meshes. The outermost layer extends beyond the margin of the compressed endocarp/seed body and is irregularly verrucate, with its surface patterned by equiaxial cell outlines c. 20 µm. Immediately inside is the layer with veins, and beneath the vein layer is one or two cell layers infilled with beige secretory substance, beneath which is densely packed fibrous tissue seen only in small damaged area; full thickness unknown because specimen otherwise complete. Without obvious perianth, style or attachment scars. Venation diverges from the more pointed end. Comment: The outer layer covering these disseminules is so thin, that in this state of strong compression, it is difficult to determine whether it covered each of them independently, or if a single layer enclosed all entities forming a fleshy fruit. The endocarps or seeds are inferred to be robust probably not originally circular in cross section but elliptical. The infilling of beige secretory substance readily distinguishes this taxon from all others known from Messel. Specimen: SM.B Me 23088.

Carpolithus sp. 36 (Pl. 61a–l) Description: More or less globose head of rigid construction, 15–18 mm wide, 13–18 mm long. Peduncle thick (3–5 mm), composed of dense fibrous tissue, up to at least 20 mm long. SM.B Me 21278 and 8469 show a thickened band near the top of the peduncle, with an obvious transverse line at the broadest point, 2.5 mm below the head. Central core of head with mostly small isodiametric cells and sinuous coarse, fibro-vascular bundles radiating from the base and later diverging. Surface of head with 4–6sided compartments, 4 mm in diameter, demarcated by a thin darker layer, best seen in SM.B Me 23879. Compartments 5.5 mm in longitudinal dimension, wall fibrous in longitudinal section. In surface view the margins of each compartment have five (rarely four) wrinkled to verrucate scale-like lobes covered with a shiny cuticle. These lobes and underlying seeds give the entire fruiting head a coarsely verrucate surface. Between these lobes and the seeds is a second set of less distinct cuticular? lobes (SM. B Me 23879). Two seeds per compartment. Seeds elliptical in outline, 3.6–4.8 mm long c. 1.4–2.0 mm wide, seed coat layered, surface cuticle finely longitudinally striate with elongate polygonal cell outlines (SM.B Me 23879, pl. 61l) up to 230 µm in length, underlying outer seed coat layer black and shiny, relatively thick (c. 100 µm) not columnar, but cell arrangement unclear. Surface 67


striae cover entire seed except in a linear groove at one end, possibly hilar. Innermost seed coat layer a very thin cuticle, cell pattern not discernable. Internal seed cavity filled with pale brown tissue. Comments: These pedunculate heads are usually found isolated, but SM.B Me 7041 (pl. 61e) shows their attachment to apparent short shoots on an elongate axis. There is some similarity to Moraceae, e.g. Maclura Nutt. which has a similar external appearance to its heads, but we have not found comparable seeds in that family. This species appears likely related to Hamamelidaceae, or Altingiaceae, both of which can form globose heads of united woody capsules and seeds with shiny longitudinally striate surface. The tentatively inferred hilar condition conforms to Hamamelidoideae. Seeds of similar size and with similar surface cell pattern to in situ are found dispersed at Messel (e.g. SM.B Me10599). These specimens recall in some ways the Messel fruits that we have assigned to Steinhauera subglobosa (Altingiaceae). This taxon may be the same in a different ontogenic state but the 4–5 lobes per compartment may be distinctive and the pedicels tend to be longer in Carpolithus sp. 36 and hence the two taxa have been kept separate here. Specimens: SM.B Me 5600 (shows seeds clearly), 5601 (striated seed surfaces visible in outer third of head), 7041, 7050, 7619, 7651, 8469, 18953, 21278, 23879, 24615; Isolated seeds, e.g. SM.B Me 10596, 10599.

Carpolithus sp. 37 (Pl. 62a–i) Description: Fruit globose to pyriform, length 20.7 mm; width 17.3–18.5 mm (n = 3). Unilocular, relatively smooth surface; wall 2.5–3 mm thick, composed of anticlinally arranged fibers whose ends form a fine polygonal pattern at the surface. Base protruding into a short broad extension with large circular scar. Apex lacking obvious style remnant. Centre filled with pale viscous spongy, possibly parenchymatous tissue. Comments: We have tentatively interpreted this as a single-seed fruit, but it might be instead a thick-walled seed. In the freshly cut specimen (SM.B Me 2299, pl. 62f, g) the internal viscous spongy substance was uniformly creamy white in colour; some months after cutting the contents became slightly darker. This substance which is darker in other specimens is likely representing the endosperm. None of the other fruits/seeds from Messel have these strikingly white contents. There are other specimens in the collections that might represent this taxon, but they have not been cut or broken to confirm internal structure (e.g. SM.B Me 2321). Specimens: SM.B Me 2299, 2583, 4848.

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Carpolithus sp. 38 (Pl. 63a–h) Description: Fruits globose, 6–8 mm diameter, with a single hypogynous whorl of 5 [or 6] thin subequal rounded tepals c. 1 mm wide. Entire structure clothed in unbranched elongate hairs 1.4–2 mm long, sinuous to straight. Hairs composed of longitudinally elongate cells, thus appearing striate. Granular layer of yellowish white resiniferous composition, c. 250 µm thick [measured on SM.B Me 15474] lies below the layer of hairs. Resiniferous layer composed of individual blebs that are irregular, with rounded surfaces, up to 160 µm in maximum dimension, and internally showing their composition of discrete smaller granules. Large areas of visible resin only occur where a specimen has been fractured as in SM.B Me 15474a, b and 15476a, b. Otherwise resin is only visible as paler streaks seen through the fine hairs. Central portion of fruit consisting of non resiniferous soft organic tissue with no discernable internal structure. A brown cuticular layer is situated between the resiniferous layer and the hair layer. Comments: These globose hairy structures are preserved with a distinctive underlying yellowish white, “resin” appearing substance. The hairs splay on compaction of the fruit, demonstrating that they are free from each other (pl. 63c–h). The position of persistent perianth parts (pl. 63e–h) indicates that these fruits developed from a superior ovary. The same fruit type is also known from Eckfeld, from which a more complete specimen shows that the fruits were borne on elongate pedicels in a paniculate infructescence. Epifluorescence analysis of the Eckfeld specimen indicates that the hairs are clothed in microprickles (VW, pers. obs). Although infructescences of Moraceae may have a globose form and produce latex, the presence of a perianth at base of the whole structure indicates that this is from a single flower. Specimens: SM.B Me 2092, 7261, 11311, 15474a, b, 15476a, b, 23336.

Carpolithus sp. 39 (Pl. 64a–m) Description: Fruit more or less elliptical, at least 25 mm long, and 30 mm wide, composed of at least two membranous valves or wings, with several subglobose seeds positioned near the centre. Each of the membranous valves approximately D-shaped with a convex-rounded, entire, outer margin and a more thickened, straight inner margin possibly representing the septum. Venation radiating from the straight inner margin, forming a reticulum almost uniform in thickness, of mostly one order of veins, with a fimbrial vein along the margin. Seeds rounded, 3.8–5.0 mm diameter, sometimes truncate at

one end, but lacking obvious scars or slits. Seed coat fractured and cracked due to brittle behaviour, black, smooth and somewhat shiny. By light microscopy, there is a surface pattern of very fine (c. 10 µm) polygonal isodiametric cell outlines. In some specimens the surface also shows sparse depressions which are less shiny (obvious in SM.B Me 4359 and 4350; seen also in 2636, 4358, 4374). Under SEM (SM.B Me 2288, 4359, 4366, 17448) the seed coat surface is composed of a pattern of barely distinguishable small polygons (7–10 µm in maximum diameter) the outlines of which presumably mark anticlinal cell walls. The polygons are mostly more or less equidimensional in surface view; some are symmetrical, others asymmetrical. Each polygon encloses an unornamented convex area assumed to be outer periclinal wall. At intervals (c. 150 µm) across the surface there are small open pits or apertures (pl. 64k, m), sometimes paired, c. 10–20 µm in maximum dimension), which may be stomata or hair bases or sites of former crystals. The surface polygons are arranged in rows which radiate from these pits and polygons surrounding the pits themselves are more elongate than those in the mid areas between pits. Seed coat multilayered (pl. 64g, h). Outer columnar layer composed of thick walled sclereids, c. 15–40 µm thick with almost totally thickened walls, lumina absent or small and visible only at the inner margin by SEM. Underneath the columnar layer is a 60–70 µm thick multilayered tissue composed of thick-walled isodiametric cells, probably parenchymatous. Comments: The seeds on the membranous fruit specimen (pl. 64a–c, g, j) correspond in morphology to those found isolated in the shale (pl. 64d–f, h, i, k–m). A very tiny fragment of one of the seeds from the fruit specimen removed for SEM analysis shows the same surface anatomy (pl. 64j) as the isolated seeds (pl. 64k) with a pattern of hexagonal cell outlines with scattered apertures, c. 10–20 µm in diameter. The venation and organization of this fruit, with hard globose seeds situated at one level, about half way, from the base of the fruit, is similar to Koelreuteria (Sapindaceae). However, the seed surface pattern examined by SEM on extant Koelreuteria seeds, does not match the pattern on these fossil seeds. The pattern of surface cells as observed under high magnification, with faintly domed roundedpolygonal cells, and scattered circular apertures (pl 64i–m), distinguishes this seed type from various other smooth orbicular seeds of generally similar outline in the Messel flora, including Carpolithus sp. 50 and 58. SEM reveals that the surface of the arillate seed (Carpolithus sp. 58) is smoother with more obscure surface pattern and no small open pits. Carpolithus sp. 57 differs in being more shiny and more brittle, typically smaller, and in SEM by a distinctive surface pattern of polygons enclosing multiple small pits. Such seeds can be difficult to distinguish from one another, if other


diagnostic features (such as the aril in Carpolithus sp. 58) are hidden within the shale. A careful study of the surface pattern needs to be made under high magnification, preferably with SEM. Staphylea (Staphyleaceae) has not been ruled out as a possible identification for this fossil. The association of smooth rounded seeds with a papery fruit is found in Staphylea. Staphylea seeds have a prominent easily recognised hilar scar but we have not been able to recognise this scar in the attached or isolated seeds. We have not examined extant Staphylea seeds by SEM. Specimens: SM.B Me 2288 (seed SEM, Collinson Frankfurt Messel stub 84), 4366 (SEM, Collinson Frankfurt Messel stub 83), SM.B Me 17448 (seeds on winged fruit) Included seed material (anatomy not studied by SEM): SM.B Me 2285, 2636, 2662, 4148, 4350, 4358, 4359, 4374, 4487, 4709.

Carpolithus sp. 40 (Pl. 65a–c) Description: Oppositely branched woody axis (5 mm thick) bearing ellipsoidal “bodies” 8–12 mm long, 4.2– 6.2 mm wide. Lower pair of “bodies” terminating equal side branches (or pedicels?), upper pairs more or less sessile. Individual “bodies” longitudinally fibrous and consisting of multiple dense layers? Comments: If the ellipsoidal “bodies” represent fruits, there are insufficient characters visible to comment on the systematic affinity, and galls can not be excluded. Specimen: SM.B Me 8956.

Carpolithus sp. 41 (Pl. 65d–h) Description: Group of hard elliptical seeds within a berry-like fruit about 12 mm diameter. Seeds ellipsoidal (length 3.8 mm, width 2–2.3 mm), with outer columnar sclerotesta (c. 100 µm thick) and longitudinally striate, shiny, inner cuticular layer. 10 seeds are visible, up to 15 inferred in cross sectional area seen. Comments: The striated appearance of the seeds somewhat resembles that in seeds of Carpolithus sp. 35, however, in Carpolithus sp. 41 this is the cuticular inner seed coat layer that is visible, not the same striate layer as in Carpolithus sp. 35. In addition, the entire structure of Carpolithus sp. 41 is much less woody with only a thin organic layer of fruit tissues on the oil-shale matrix. It appears that some of the structure of this specimen may still be embedded in the oil shale beneath the part illustrated. The counterpart shows almost no material. Specimen: SM.B Me 2328.

69


Carpolithus sp. 42 (Pl. 66a–p) Description: Fruit with granular to smooth surface, 10.3– 5.9 mm long and 7.6–3.6 mm wide, ovoid with rounded apex and truncate base with an elliptical indentation marking the attachment. Outermost fruit wall rarely preserved, very thin (c. 100 µm, at base in SM.B Me 21364, pl. 66f, at edges in SM.B Me 21337), outermost part lacks obvious structure, innermost part consisting of rectangular cells with dark brown convex outer periclinal walls arranged in longitudinal rows giving a striate appearance (SM.B Me 21366, pl. 66c). The underlying, and usually most obvious, fruit wall layer (0.25–0.6 mm thick, n = 8) is black, shiny and amorphous, fracturing with blocky to conchoidal fracture into irregular typically cuboidal (to polygonal) pieces (pl. 66b, i, k). In most specimens this is the outermost layer preserved, it is almost always fractured and portions of the layer are often broken away. Underlying the amorphous blocky black shiny layer there are a few black fibres in a discontinuous layer (SM.B Me 21365). A fortuitous split during preparation (SM.B Me 21350, pl. 66d) has released an inner fusiform structure with rounded apex and slightly truncate base consisting of inner fruit wall tissues. Longitudinal fibre strands (up to 8 estimated in total) underlie the blocky layer, one in the median line is more prominent on the inner wall and is represented by a groove on the inside of the black shiny blocky layer (median fibre strand also seen in SM.B Me 21395 and 5017). These fibre strands may represent vascular tissue. The inner fruit wall layer is dark brown and lacks obvious structure in LM. The innermost layer seen lining the locule is cellular and darker brown. In SEM the black shiny blocky layer seems initially amorphous but at high magnification shows a remnant cellular organization (pl. 66n). Cells are equidimensional and completely infilled with no residual lumina. The infill is granular possibly suggesting a secretory origin (pl. 66l). Occasional tiny cavities remain at the corners of cell junctions. The layer beneath the black shiny blocky layer is cellular, most cells are infilled with infill of similar texture to the black layer. In some places this layer is fibrous and in others the cells are more equiaxial (pl.66m, o, p). The tissue lining the locule is variable, perhaps due to slightly different compaction or to position in locule, or both. It is composed of equidimensional cells with open lumina or with closed lumina (pl. 66p) or of platy cells (pl. 66o). The locule is lined by a cuticle with elongate cell pattern. Comments: This taxon is distinguished by the amorphous shiny black layer of the fruit wall which exhibits blocky to conchoidal fracture and resembles a bright coal in appearance. No other Messel fruit or seed shows this type of organic material as a wall layer. It seems likely that this layer originally possessed a distinctive chemistry perhaps a secretory tissue or a tannin/polysaccharide 70

composition. Van Bergen et al. (1997) argued that fruits/ seeds with tannin/polysaccharide thickened walls would be lacking in the typical fossil record but the exceptional preservation at Messel could have resulted in their survival. The botanical affinity of these fruits is unknown. Specimens: SM.B Me 2330, 2594, 2601, 2602, 2613, 4434, 4495, 4640, 4643, 4644, 4654, 4659, 4665–4667, 4685, 4694, 4695, 4697, 4708, 4752, 4774, 5008–5034, 5581, 5647–5661, 5663–5676 (including 5664 sectioned longitudinally), 5678–5708, (5658, 5690 & 5703 all sectioned transversely), 5740, 21281, 21286, 21297, 21308, 21314, 21315, 21319, 21327, 21328, 21333, 21334, 21337, 21350, 21352 (rare example showing very little blocky fracture), 21359–62, 21364–67, 21372, 21374, 21378, 21379, 21389, 21390, 21395, 21396, 21398– 21400, 21403, 21405. Probably this taxon 5034–5045 (coaly layer not clearly revealed) and 5709–5722 (unprepared).

Carpolithus sp. 43 (Pl. 67a–d) Description: Fruit very large, elliptical (92 mm long, 60 mm wide in SM.B Me 7060), highly compressed, surface of the broken interior appears multifragmented showing some light patches of different shape, sometimes with a finely pitted surface. Surface of the fruit preserved in a few small areas, obviously fibrous. Several elliptical seeds are seen within the central region of the fruit. Seed elliptical to ovate, 7.5 mm long, 5 mm wide. Total number of seeds, their arrangement and placentation uncertain. Comments: These are the largest fruits known from Messel. The fractured surface preserved in the two specimens studied, suggests deformation of a fruit with a brittle outer layer. The internal tissue was apparently relatively soft because the fruit is very strongly compressed in the shale. Although flattened elliptical seeds can be seen (pl. 67b, c), details of placentation and internal seed anatomy remain unknown. Specimens: SM.B Me 7060, 7061 (fragmentary).

Carpolithus sp. 44 (Pl. 67e–g) Description: Fruits fusiform, borne on a twig with two apparently opposite leaves and a third leaf, the petiole of which is attached slightly higher. Fruits not necessarily axile, but appear to be borne on the petiole. [an alternative interpretation is that the leaves are sessile, with the apparent petioles being instead short twigs]. Four fruits preserved (one partially hidden); (1 or 2 fruits/leaf observed), fruits (sub)sessile, elongate, 10–13 mm long and 2.6–2.8 mm wide, with longitudinal grooves. Distally, the fruits are constricted and tapering above into an apical beak-like structure, 2.5–3 mm long. Perianth scars not obvious to indicate whether inferior or superior. The at-


tached leaves are entire-margined and pinnately veined with a marginal vein. Twig naked, 13.5 mm long (no scars along that length). Tertiary veins obscure. An ellipsoidal body apparently glandular, positioned immediately below each lamina, 1 mm long by 0.6 mm wide with long axis parallel to the leaf axis. Comments: This unique specimen is highly unusual in preserving attached leaves and fruits. We are not aware of isolated fruits of this kind in the assemblage. If the phyllotaxy is correctly interpreted as opposite, then the families Oleaceae, and Combretaceae would be appropriate candidates as some of their extant genera have fusiform fruits. The possibility remains however that these could be flower buds or galls. There is no evidence of stamens or pollen. The ellipsoidal bodies below the laminae (pl. 67g) might be glands, insect eggs or galls.

either side, with transverse striations, and partial preservation of at least one additional but less pronounced similar ridge. Fruit wall not thick, very collapsed giving little direct evidence for the nature of internal tissues; cuticle preserved all around the fruit. Seed morphology unclear, at least 2 mm in longest dimension, indicated by remnants of dark seed coat near the apex of the structure, showing polygonal cellular structure inside the surface [outer surface of seed coat obscured]. Comments: This fruit is distinguished by large size, strikingly stout pedicel, hypogynous perianth, and the fleshy or weak tissue making up the main body. Although fragments of only a few seeds are preserved, the size of the fruit indicates that they must have been numerous. Specimen: SM.B Me 7073.


Carpolithus sp. 47 (Pl. 68e–h) Carpolithus sp. 45 (Pl. 67h–k) Description: Crescent-shape fruit 24–29 mm long, 13.5– 15 mm wide, with 5 valves, 3 of which are visible. One of the specimens (pl. 67j), shows a median groove on the dorsal surface of the valve but this was not observed in the other, (pl. 67h). Fruit narrowing to both ends, with a truncate base with a large circular attachment scar c. 3–4 mm diameter; shed without pedicel. Apex, terminating in a 5-lobed protruding pad of tissue (stigmatic?) 0.8 mm in diameter. Strong longitudinal ribs at the valve junctions. Valves sometimes (SM.B Me 20418) with discontinuous and anastomosing finer ribs. In SM.B Me 25000 surface smooth, without obvious cellular pattern. Internal structure unknown. Comments: These banana-shaped specimens are inferred to be dicotyledonous with five carpels, based on the 5lobed apical protrusion/pad at apex (pl. 67i). This taxon lacks the surface hairs present on similarly shaped fruits of Carpolithus sp. 3. Specimens: SM.B Me 20418, 25000.

Carpolithus sp. 46 (Pl. 68a–d) Description: Large globose fruit, 34 mm diameter, on a thick pedicel (9 mm thick, at least 11 mm long) with a hypogynous perianth of more than 7 parts partly overlapping each other but not showing clearly whether organised in one or two whorls, the exposed tepals wider than long. The fruit has a prominent longitudinal ridge with a median groove at the crest with dark band of tissue on

Description: Specimen broadly ellipsoid, 3.8 mm long, 3.0 × 2.0 mm wide, somewhat D-shaped in cross section, rounded at one end, pointed at the other. Ornamentation consisting of c. 18 rounded longitudinal ridges that extend to both ends of the object. Bilaterally symmetrical with a more prominent ridge on the more rounded side in the plane of symmetry, and with a groove on the opposite (flatter) side in the plane of symmetry. Comments: Despite the similarity in size and ridging, this specimen differs from that of Spirellea in shape in lateral and face view and by the more rounded ridges lacking transverse striations. Specimen: SM.B Me 7302.

Carpolithus sp. 48 (Pl. 68i) Description: Fruit widely ovate, 2.6 mm high, 2.2 mm wide, with a median longitudinal rib (possibly a keel) in face view, apparently having been at least trigonal, with another possible keel exposed at the lower left margin. Surface with a longitudinal cellular pattern. Two apical protrusions appear to represent styles. Comments: Although known from only a single specimen, this fruit is distinguished by what appears likely to have been trigonal symmetry with three equal faces, two of which are facing upward separated by a longitudinal rib (pl. 68i). Two stylar arms are attached, and another, which is no longer attached lies horizontally near the apex of the fruit. If this was a trigonal fruit with 3 style arms, then it could be Fagaceae, or Polygonaceae? or Cy71


peraceae. The position of perianth, whether epigynous or hypogynous, is not clear from this specimen, however. Specimen: SM.B Me 23185.

Carpolithus sp. 49 (Pl. 69a–f) Description: Infructescence with woody axis, c. 1.5 mm thick bearing subsessile fruits either oppositely or in whorls of 3–?4. Internodes about 5 mm in length. Fruits obovoid with persistent calyx extending 1/3 to 1/2 the length of the fruit, of which parts of three lobes can be seen in face view (estimated 5 in total from symmetry) (SM.B Me 8950, pl. 69d–f). Fruits 5.5–9.5 mm long, 5–6 mm wide, splitting longitudinally into 5 persistent valve like units. The calyx lobes seem to have been shed at maturity leaving behind a cup-like structure covering the lower one quarter of the fruit (SM.B Me 7621). Stylar area unclear, seeds not preserved. Comments: This infructescence, with its woody axis and sessile fruits, is similar in gross form and size to those of Hamamelidoideae such as Corlyopsis maii (pl. 14a–e), but is readily distinguished by the lack of persistent styles and has five valves (pl. 69f), rather than the two seen in Hamamelidaceae. The persistent perianth lobes are well seen in one of the specimens (pl. 69b). Although similar in some respects to the capsules of Theaceae and Ericaceae, these fruits do not appear to retain a central column on dehiscence. Specimens: SM.B Me 7621, 8950.

Carpolithus sp. 50 (Pl. 69g, h) Description: Small operculate seed, widely ellipsoidal, length 7.3 mm, breadth 6.5 mm, probably originally spherical as indicated by radial and longitudinal fold and cracks, compressed, not brittle and not fully flattened suggesting some thickness, ductility and strength to the seed coat. Operculum surrounded by raised rim, the margin of which is sharply demarcated from the operculum area. Operculum 1.65 mm long and 1.8 mm wide, surface dull and rough with no discernable cell pattern. Centre of operculum with small columnar protrusion c. 0.5 mm diameter and 0.25 mm high. Seed coat as seen in LM only, in places (rim, left edges) smooth, shiny and structureless, in places (left base) dull, slightly corroded giving a digitate pattern. In other areas (right edge) seed coat with raised narrow ridges sometimes forming a poorly defined reticulum (lumina 30–40 µm in maximum dimension). Comments: The single specimen has not been dissected for SEM or used for SRXTM study. Taxonomic affinity 72

to Sargentodoxa Rehder & E. H. Wilson is supported by the distinctive operculate morphology (Tiffney 1993: fig. 82H). The fossils described by Tiffney (1993) all lacked the operculum and if compressed as in Messel oil shale the former presence of the operculum could certainly have been obscured so that additional specimens from Messel might remain unidentified. Although superficially similar to seeds of Carpolithus sp. 58 detached from their arils, this species is distinguished by: (1) a much smaller operculum (less than one third of seed diameter) (2) a mostly dull seed coat with no obvious small polygonal surface cell pattern and (3) the nature of compaction (flattened) and damage (elongate crumple) which implies a ductile not brittle seed coat. There is also no evidence for the former presence of an aril on this single specimen. Specimen: SM.B Me 4791.

Carpolithus sp. 51 (Pl. 70a–f) Description: Seed subovoid 3.9–4.3 mm longest dimension, 3.2–3.6 mm in shorter dimension (n = 3), patches of a thin leathery shiny outer cuticle with a surface pattern of fine polygons c. 20 µm, diameter survive on the outside of the seeds. Underneath this layer is another layer the surface of which appears slightly striate due to very fine rugulae or verrucae. Single-layered columnar sclerotesta 0.2 mm thick expanding to reach 0.5 mm in thickness in a mound at one end which is 1.4 mm diameter. There is a narrow opening through the centre of the mound, which expands inwards forming a vestibule. Within the seed there is a thin cuticular layer with a polygonal pattern discernable under the mound. Comment: Co-occurrence of two seeds on one specimen suggests they were from a single multiseeded fruit, or from a coprolite. Although only preserved in patches, the outer cuticle appears to have covered the entire seed including the mound, and is also crumpled in places, suggesting possible presence of a softer tissue, which has now decayed, originally between the cuticle and the columnar sclerotesta. In all three specimens the end opposite the mound is not compressed, suggesting a very robust and thicker sclerotesta in this region. Specimens: SM.B Me 21394, 21397 (two specimens on one slab).

Carpolithus sp. 52 (Pl. 70g–i) Description: Disseminule elliptical in outline, 5.2 mm wide, 8.1 mm long, probably originally broadly lenticular in cross section, broken at one end with two lobes


protruding beyond the broken wall. At other end a small indentation (0.8 mm diameter). Longitudinally gently ribbed with low topography, these ribs 1.0–1.5 mm wide. Outermost surface layer formed of cells that are mostly square to polygonal-equiaxial in surface view, 20 µm diameter. Broken wall exposes compacted tissue that does not appear to have been woody, columnar, or fibrous. The protruding lobes are a discrete inner tissue layer, lacking an obvious surface cell pattern. Comment: The two lobes protruding from the broken end of this structure might represent a pair of seeds, or a pair of cotyledons within one seed. Physical and/or digital dissection of this specimen is needed to reveal additional details of internal morphology and anatomy.

lain by non columnar, possibly isodiametric cells. External surface of fruit lacking obvious ornamentation, fibrous. Comments: This large, bilocular, presumably bicarpellate woody fruit is represented by a single specimen exposed in ventral view. The three-dimensional symmetry indicates that this represents half of the original fruit. However, there is no smooth surface to indicate that it fractured along a natural dehiscence plane. The bulbousness of the locules accommodated seeds that would have been approximately circular in cross section. There is a superficial similarity in shape and size to nuts of Carya and Juglans (Juglandaceae), however the locules are fully separated by the central septum rather than having a single seed with partial septum of Juglandaceae, and there is no obvious plane of dehiscence.

Specimen: SM.B Me 21575. Specimen: SM.B Me 23874.

Carpolithus sp. 53 (Pl. 71a–f) Description: More or less circular, originally very soft, fleshy fruit containing clumps of flat elliptical seeds. Fragmentary yellow cuticle preserved, enclosing the entire structure, probably fruit epidermis. No pedicel or style. Fruit 10.9–12 mm diameter. Long axes of seeds aligned radially in the fruits. Seeds with a finely pitted black outer layer, and internal cuticle, without visible raphe, chalaza, or hilum, internal cuticle of the seed elliptical 2.0–2.2 mm long and about 1.8 mm wide. Comments: Judging from the pronounced flattening, the thinness of tissues, and the faint pigmentation, this is one of the fleshiest, wateriest fruits of the Messel assemblage. We consider these to represent fruits rather than coprolites, because of the enclosing cuticle and radial alignment of the seeds. Specimens: Darmstadt example: HLMD-Me-9078 (Heil et al. 1987, fig. E6), SM.B Me 7071 (seed shows only inner cuticle), 7353, 10523.

Carpolithus sp. 55 (Pl. 72a, b) Description: Four nutlets/pyrenes situated in the centre of a circular wing. Wing 18 by 21 mm in diameter, membranous, with fine dichotomizing and anastomosing venation. Pyrenes closely adjoining each other in the centre of wing, forming a collective central body 9–10 mm wide. Each pyrene rounded dorsally, with a pair of flattened ventral faces forming a 90° angle at the mutual junction with the other three pyrenes. Surface mostly smooth but with wrinkles 5–5.5 mm dorsiventral dimension, 5.0 mm wide. Comment: This fruit has organization similar to Paliurus Tourn. ex Mill. (Rhamnaceae) and Cylcocarya Iljinsk. (Juglandaceae) which have a thin circular wing at right angles to the axis of symmetry, with a central nut. However, in the Messel specimen there are four separate symmetrical pyrenes or nutlets in the centre. The affinities remain uncertain. Specimen: SM.B Me 25013.

Carpolithus sp. 54 (Pl. 71g–i) Description: Large woody fruit, cordate in face view, 28 mm long, 36 mm wide. Fruit wall thick (2–3 mm) composed of longitudinal fibers, septum also 3 mm thick, composed of longitudinal fibers. Pointed end broken, opposite end with bulge in the cleavage area but broken at the extreme tip beyond the bulge. Locules two, D shaped, attenuate apically, 9 mm wide and 23 mm long. Locule lining black and shiny in fragmentary preservation, about 40 µm thick, in places striated, firmly fixed to the fruit wall. Remnants of possible seed with black shiny brittle outer layer very thin c. 10–15 µm thick, nearly smooth, under-

Carpolithus sp. 56 (Pl. 72c–h) Description: Seed circular in outline 1.5–1.8 mm diameter, and flattened in the plane of compaction. Surface smooth. An elongate groove is seen along edge of one of the specimens, possibly a raphe. External surface finely pitted, resulting from an underlying probably polygonal cellular pattern. Comments: These small flattened orbicular seeds would easily be overlooked during collecting from bedding 73


planes. Thus, they may be under-represented in the Messel collections compared to fruit and seed floras collected by the sieving method. Specimens: SM.B Me 4132, 4138, 4533, 15115 (multiple seeds on same shale specimen). SM.B Me 4563 is also small and has the same pitting, but is more inflated and angular, with visible scars so it may represent a different species.

Carpolithus sp. 57 (Pl. 73a–o) Description: Seed globose to ellipsoidal, 3.4 × 2.8 mm, 3.8 × 3.6 mm, to 4.25 × 3.1 mm (n = 6), frequently distorted, shattered or incomplete, mostly lacking any obvious scars or slits. In one case (SM.B Me 4012) there is a circular scar (possible hilum) at one end and slight lateral faceting either side of a longitudinal ridge which might indicate position of a raphe. Seed coat fractured, cracked and shattered due to brittle behavior, black, smooth and very shiny. With appropriate side lighting, a surface pattern of tiny polygons can be seen (pl. 73f). SEM reveals a distinctive seed coat anatomy the outer surface of which has a polygonal (typically hexagonal) pattern. Each polygon is formed from rounded ridges (presumed outer ends of anticlinal cell walls) which enclose a flat area containing from 10–13 small pits. The pits are closed by the covering of the outer periclinal cell wall. The polygons are mostly more or less equidimensional and vary from c.18–25 µm in maximum dimension, some are symmetrical and others asymmetrical. The small pits are more or less equally spaced and typically 3–4 µm in diameter. The pits are arranged all around the inner edge of the polygon ridges with three or four in the central area of the polygon. SM.B Me 4383 and 5738 show clearly the characteristic polygonal pattern and very shiny surface of this taxon. In addition, a small area on their surfaces has a pattern of fine parallel striations (pl. 73g, h). Two sets of parallel striations diverge at a low angle in two directions from a mid line (pl. 73g). This structure is rarely seen (presumably in most cases it is still obscured in the oil shale or has been lost due to damage on these very brittle specimens). However, a further four specimens (SM.B Me 2161, 4379, 4380 and 4875) also show striations though not details of their precise organisation. The number of specimens and the clear organisation seen (especially in SM.B Me 4383 and 5738) confirm that these striations are an original feature of the seeds. When dissected the seed coat separates into two layers. The outer layer is a sclerotesta c. 70 µm thick with several layers of polygonal cells with almost totally thickened walls, the outermost possessing very clear lumina just underneath the outer surface (pl. 73o). These lumina correspond in size to the small surface pits and possibly represent the former position of crystals in the seed coat. The inner surface of the outer seed coat layer has a polygonal cell pattern comparable to that of the 74

outer surface of the inner seed coat layer. The inner seed coat layer has an outer part, several cells in thickness, consisting of polygonal more or less equidimensional but radially flattened parenchyma cells (pl. 73k, m, n). The central part consists of elongate thick-walled fibrelike cells with at least two orientations perpendicular to one another (pl. 73k, m, n). The innermost layer is a thin cuticle with outlines of elongate epidermal cells arranged in rows (pl.73l). Comments: These globose to ellipsoidal seeds are black, shiny, and commonly fractured (pl. 73a–e). Under reflected light microscopy, a striate appearance may sometimes be detected from aligned cells (pl. 73e, g, h). Higher magnification from SEM reveals a distinctive pattern not known among other Messel taxa: a surface consisting of polygons outlined by rounded ridges and enclosing c. 12 small pits (pl. 73i, j). Smooth black shiny seeds of various extant plants were examined for comparison with these fossils including Dillenia L.; legumes such as Erythrina L., Euphorbia L., Illicium L., and Sargentodoxa but nothing has been seen with this distinctive seed coat surface anatomy. We considered whether the Carpolithus sp. 57 fossils might be a specific preservation or maturity state of a taxon where the surface pattern normally appeared simpler. This possibility can be rejected for several reasons: (1) The pattern is consistent in four different seeds examined by SEM and in the numerous fragments found filling the gut content of one specimen of the rodent Masillamys (Collinson 1988). This would be very unlikely if the pattern were a consequence of preservational conditions, (2) where the seed surface can be seen free of sediment the anatomy is seen to be uniform; it is not patchy as might be expected if affected during preservation/digestion, (3) small ‘lumina’, possibly former crystal sites, are present just beneath the surface (seen in vertical section) apparently one corresponding to each small pit. Such features are absent in other smooth black shiny seeds from Messel studied by SEM and from comparable modern seeds. (4) It is hard to understand the large numbers of dispersed seeds if they are immature. Specimens: Studied in detail: SM.B Me 2162, 2287 (dissected), 2293, 2631 (MEC stub 2242), 4012, 4031 (MEC stub 2244, now dissected), 4083, 4346 (MEC stub 2251; Collinson Frankfurt Messel stub 87), 4354, 4365 (Collinson Frankfurt Messel stub 85), 4371 (Collinson Frankfurt Messel stub 86), 4375, 4383, 4721, 4875, 5738. Other referred specimens: SM.B Me 288, 308, 309, 2160, 2161, 2284, 2286, 2287, 2290–93, 2622, 2648, 2650, 4119, 4343, 4348, 4351–53, 4356, 4360–65, 4367, 4370, 4372 (with boring?), 4376, 4377–4382, 4384, 4385, 4464, 4528, 4542, 4612, 4662, 4668, 4690, 4716, 4729, 4734, 5007, 5738, 23119.

Carpolithus sp. 58 (Pl. 74a–p) Description: Seed subglobose, length including aril 7.7–9.8 to 11.1 mm (length of specimens lacking aril

5.2–8.0 mm), width 5.1–6.7 mm, smooth, shiny, arillate and transversely truncate beneath the aril. Aril substantial, compact, caplike, affixed to the truncate end (considered to be the hilar end), symmetrical, 3.2–5.9 mm wide, 1.8–4.4 mm high, composed of fibers radiating from attachment (pl. 74h). Truncate end with clearly defined dull area beneath aril, dull area mucronate with central protrusion, sometimes with a small hole (SM. B Me 4731). SM.B Me 11770 (pl. 74k) shows a more extended protrusion than the other specimens. Seed coat surface almost smooth even by SEM, with poorly defined polygonal cell outlines (pl. 74o), 8–10 µm in diameter. Seed coat multilayered, totalling at least 250 µm in thickness. The outer layer c. 160 µm thick, composed of anticlinally aligned rows of sclereids, giving a columnar appearance (pl. 74l–n); SEM reveals scattered small angular cavities – rectangular to diamond-shaped, to rounded (idioblasts?), 0.8–3.2 µm within this layer. The next underlying layer c. 30 µm thick, composed of thick walled isodiametric sclereids with almost no lumina (pl. 74n). The next underlying layer is composed of loosely adhering tabular cells and is about 30 µm thick (pl. 74n). Underlying this is a tissue c. 25 µm thick with several layers of blocky bricklike cells, closely packed with no discernable lumina (pl. 74n). Innermost layer is a cuticle lining (pl. 74p) with a cellular pattern indicating underlying elongate epidermal cells arranged in rows, cells c. 70 µm long, 7 µm wide. Raphe not obvious on seed surface. Comments: These seeds are distinctive by their smooth surface and circular truncation on which is seated a prominent domed aril (pl. 74a–d). When detached from the aril (pl. 74e–g), these seeds might at first be confused with those of the previously described taxon, Carpolithus sp. 57 (pl. 73). Although both types are smooth and shiny, the seeds of Carpolithus sp. 58 (pl. 74) lack the surface pattern of polygons each containing c. 12 small pits. The well-marked circular mucronate truncation in the seed underlying the aril (pl. 74 g, i, j, k) might represent an operculum, but we have not seen any specimens with an opening or space in this position, nor confirmed it by sectioning. Among the arillate seeds we have considered, some Zingiberales have a much smaller operculum in relation to seed diameter and a less compact aril (cf. Manchester & Kress 1993: 1268–1269). Smooth seeds with an aril of coarse hair-like filaments occur in the three extant genera of Strelitziaceae but the hair like filaments are often much less compact in arrangement. In overall morphology, this seed greatly resembles that of Phenakospermum Endl. (Strelitziaceae), of tropical America (Cornejo & Janovec 2010: p. 139), however, the fossil has a wide, flat truncation apically beneath the aril, that is lacking in Phenakospermum. Seeds of Sapindaceae can also be smooth and shiny, and may be arillate, but the aril is not known to be so substantial.


Specimens: SM.B Me 12, 2279–2281 (2280 with patch of aril at left), 4387, 5752, 7195, 11770, 14807 (patch of aril on clear operculum, aril also seen at left and around edge of operculum in small residual patches). Without aril: SM.B Me 7146, 4731.

Carpolithus sp. 59 (Pl. 75a–d) Description: Fruits subglobose 10.0–11.5 mm in diameter, pedicel 2.5–7 mm long, borne on an elongate twig; fruit with a prominent, coarsely longitudinally wrinkled cup-like structure preserved over the lower half. Upper half of fruit shows in face view 2 meridional ribs and parts of three apparent valves, with symmetry suggesting a total of 5 carpels. Comments: The cup like structure possibly representing a hypanthium distinguishes this fruit type from most of the others known from Messel. One of the specimens (pl. 75a, c) shows one apparently complete fruit. A fragment of a second fruit occurs on the same slab (upper left in pl. 75a) in a position that might correspond with an attachment to the same twig. Another specimen (pl. 75b, d) shows the same morphology indicating that a relatively unlobed rim of the cuplike structure is a uniform feature of this taxon and not an artifact of preparation. The prominence and position of the longitudinal ribs, suggests that this fruit may have been a capsule with longitudinal dehiscence from the apex, but we have not observed dehiscent specimens and do not know the nature of seeds and placentation. Specimens: SM.B Me 2573, 8803.

Carpolithus sp. 60 (Pl. 75e, f) Description: Fruit ellipsoidal, 25 mm long, 17.5 mm wide, surface densely verrucate to rugulate, thin and laterally cracked due to compaction. There is no obvious scar of attachment. Closely packed irregularly shaped structures (seeds?) barrel-shaped to ovoid but none fully visible (visible diameter 1.2 mm). One of the seeds has a circular truncation at one end but it has no central prominence. Comments: This fruit is distinctive owing to the numerous closely packed small irregular bodies interpreted as seeds, and thin, apparently leathery, surface (pl. 75f). There is no obvious indication of attachment scar on the fruit. The seeds of this taxon should not be confused with those of Carpolithus sp. 30 (pl. 58h, i) because they are smaller, not as shiny, and more irregular in shape. However, there is no clear view of the testa so one cannot tell if the surface is really shiny. The fruit wall is thin and 75


cracked indicating deformation of an originally ellipsoidal fruit.

Genus Equisetum L. Equisetum sp. (Pl. 76 e, f)


Carpolithus sp. 61 (Pl. 75g) Description: Globose fruit, 9 mm long, 9.5 mm wide, with a hypogynous stout (?woody) disk (width 4 mm), remaining on a branched infructescence. Pedicel 6 mm long, 1 mm wide. Fruit radially septate with at least four segments exposed on fractured surface. Remains of style(s) or seed(s) not preserved in the single specimen. Comments: Among the other branched infructescences known from Messel (Decodon, sp., Carpolithus sp. 12, 16, 17, 29), this one is distinguished by the combination of globose shape, and prominence of the basal disk. If the disk represents the site of attachment for perianth parts, this would have been derived from a hypogynous flower. The single specimen is preserved with the fruit fractured longitudinally, showing internal morphology, but the external ornamentation (e.g. whether striate or not) is not exposed. Specimen: SM.B Me 8697.

Carpolithus sp. 62 (Pl. 76a–d) Description: Fruit with a pair of prominent wings arising near the base of wide-ovoid gynoecium. Fruit body 7 mm in diameter and at least 1.7 mm thick, with a rugose outer surface and thick short ridges running from the base to apex of the fruit. The apical part of the gynoecium, between two wings, is covered in c. 13 scale-like plates, somewhat rhomboidal, arranged in c. 5 spirals. Each plate is c. 340 × 295 µm in diameter. Resin globs possibly produced by this fruit occur just above these plates at the gynoecium apex. The two wings are attached at the base of fruit and directed apically but crumpled/folded across the gynoecium. Wings narrow, elongate, entire-margined c. 27 mm long, broader in the middle and tapering to rounded apices. Wings not papery, at least c. 110 µm thick. Details of wing venation are not preserved, wing surface is covered by a wrinkled cuticle. Comments: This fruit has similarities in organisation to the biwinged fruits of Dipterocarpaceae, however, venation of the wings, which is distinctive for winged fruits of that family, and other families having convergent winged fruit morphology (Anacardiaceae, Hernandiaceae, Juglandaceae) cannot be discerned. Specimen: SM.B Me 20618.

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Description: Disseminule almost globose, 10 mm in diameter, complete, fully three dimensional (not very compressed) with surface striations formed by aligned quadrangular surface cells and a ring of several (?6 or 7) protrusions surrounding a scar (probably attachment) at one end. Comments: This specimen corresponds closely to those Manchester (1994) named Striatispermum Manchester from the Eocene Clarno Formation. Subsequently Manchester (unpublished) has determined that those specimens represent not seeds, but Equisetum L. tubers. Fritel (1910: 20, textfig. 8, pl. 1, figs 15, 16) illustrated anatomically preserved specimens from the Marnes calcaires subordonnees aux lignites de Bretigny et de Murancourt (Oise), Sparnacien of the Paris Basin. Łańcucka-Środoniowa 1969 illustrated modern and Miocene specimens including a sketch of the broader end of one of them showing 8 triangular lobes and a small central protrusion ('sheath and beak') very similar to the Messel specimen. A number of similar specimens, some in connection with rhizomes, from the Wealden (Cretaceous) of England are housed in NHMUK (Collinson personal observations). Specimen: SM.B Me 4190 (loose).

Acknowledgements This work is dedicated to the late F riedemann Schaarschmidt who realized the potential of the palaeoflora from Messel, initiated and stimulated modern research, and introduced the authors to the material. Technical help and assistance in many ways was always provided in a generous and friendly manner by Karin Schmidt and Martin Müller at the Senckenberg Forschungsinstitut und Naturmuseum in Frankfurt am Main (SFN), and we also appreciate their major contributions to curation and preparation of the specimens. Marisa Blume, Gabriele Gruber and Norbert Micklich repeatedly looked for specimens in the collections of the Hessisches Landesmuseum Darmstadt and kindly organized short-term loans. Taxonomic advice on particular systematic groups was kindly provided by Iju Judy Chen (Vitaceae), Fabiany Herrera (Menispermaceae), Sara Hoot (Menispermaceae), Frédérik Jacques (Menispermaceae), Dieter Mai (mastixioids), Selena Smith (monocots), and Greg Stull (Icacinaceae). Comparative work on extant fruits and seeds was supported by keepers and curators of Harvard Herbaria, Cambridge, MA, Missouri


Botanical Garden Herbarium, St Louis, MO, the fruit and seed collection of the Humboldt University, Berlin, and the National Seed Herbarium, Beltsville, MD (subsequently moved to National Arboretum, Washington, DC). Keepers and staff of the Palaeontology Department, Natural History Museum, London, NHMUK (=BMNH) provided access to comparative material in their care and loaned samples for study. Our thanks to Mike Howe and Susan Martin (British Geological Survey, Keyworth) for information on, and photography of, Vitis hookeri from Bovey Tracey, UK. We thank Federica Marone, Marco Stampanoni and other staff for enabling our work at the Swiss Light Source (SLS), Villigen, and Selena Smith, Sarah Joomun and Lauren Howard for their help with running samples at SLS; Richie Abel and Lauren Howard for enabling microCT scan work at the NHMUK; Tony Brain (formerly at Centre for Ultrastructural Imaging, King’s College, London) for help with electron microscopy; and Terry Lott, Florida Museum of Natural History, Gainesville, FL, for help in text processing and compiling the figure captions. We thank DAVID FORD who obtained some of the SRXTM images whilst undertaking a BSc project supervised by MEC. MEC gratefully acknowledges the late Friedemann Schaarschmidt for his help and hospitality during her study visits at SFN in the 1980s, and NERC (1978–1980), the Royal Society University Research Fellowship (1983–1993), and the Departments of BioSciences, Kings College London and Earth Sciences, Royal Holloway University of London for partial funding. SRM acknowledges support from the National Science Foundation (EAR 0174295, BSR 0743474), and the H. Kämpny family for accommodation and support during extended visits working at SFN. Helpful review comments were provided by Kathleen B. Pigg and Bruce H. Tiffney. Last, but not least, thanks are due to Senckenberg staff and the members of numerous fieldteams of different institutions who collected enthusiastically at Messel over the past thirty years. Some financial support for printing this monograph was provided by the Landesamt für Denkmalpflege Hessen, Wiesbaden, and is gratefully acknowledged.

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Manuscript submitted: 16.05.2011 Revised manuscript accepted: 22.10.2011

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Tables 1–3 Table 1: List of Messel fruit and seed taxa with families presented in the phylogenetic sequence of APG III (2009)1.

Conifers Family: Doliostrobaceae Doliostrobus taxiformis (Sternberg) Z. Kvaček emend. Kunzmann (Pl. 1a–c) Angiosperms Order: ?Nymphaeales Unnamed seeds (Pl. 35a–i) MAGNOLIIDS LAURALES Family: Lauraceae Laurocarpum sp. 1. (Pl. 22a–m) Laurocarpum sp. 2. (Pl. 22n) Laurocarpum sp. 3. (Pl. 21e–g) Lauraceae genus indet. 1 (Pl. 21c–d) MAGNOLIALES Family: Magnoliaceae Magnolia sp. fruit (Pl. 27g) Magnolia spp. seeds (Pl. 27a–f) Family: Myristicaceae Myristicacarpum sp. (Pl. 34o, p) MONOCOTS PANDANALES Family: Cyclanthaceae Cyclanthus messelensis S. Y. Smith, Collinson & Rudall (Pl. 8a–e) ARECALES Family: Arecaceae Friedemannia messelensis gen. et sp. n. (Pl. 5a–n) POALES Family: Cyperaceae Volkeria messelense Smith, Collinson, Simpson, Rudall, Marone & Stampanoni (Pl. 8f–i) EUDICOTS Family: Sabiaceae Meliosma sp. (Pl. 38a–f) RANUNCULALES Family: Menispermaceae Diploclisia rugulosa sp. n. (Pl. 31a–d) Stephania hootae sp. n. (Pl. 31e–h) indet. Menispermeae (Pl. 31i) Martinmuellera tuberculata gen. et sp. n. (Pl. 31j–m) Palaeosinomenium ornamentum sp. n. (Pl. 32b–d) Palaeosinomenium venablesii Chandler (Pl. 32e) Palaeosinomenium sp. (Pl. 32f–i) ?Pericampylus sp. (Pl. 32j) unnamed Menispermaceae (Pl. 34k) Wardensheppeya sp. (Pl. 32a) Cocculus lottii sp. n. (Pl. 32k–r) Karinschmidtia rotulae gen. et sp. n. (Pl. 33a–o) Tinomiscoidea jacquesii sp. n. (Pl. 34a–c) Parabaena cf. europaea Czeczott & Skirgiełło (Pl. 34d–f) Tinosporeae sp. 1 (Pl. 34g, h) Tinosporeae sp. 2 (Pl. 34i, j) Tinosporeae sp. 3 (Pl. 34l–n) CORE EUDICOTS SAXIFRAGALES Family: Altingiaceae

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Steinhauera subglobosa Presl emend. Mai (Pl. 1e, f) Family: Hamamaelidaceae Mytilaria boglei sp. n. (Pl. 12, 13a–f) Corylopsis maii sp. n. (Pl. 14a–e) Corylopsis waltheri sp. n. (Pl. 14f–t) ROSIDS VITALES Family: Vitaceae Vitis messelensis sp. n. (Pl. 42a–h, l). Parthenocissus britannica (Heer) Chandler (Pl. 42i–k, m–r) Ampelopsis sp. (Pl. 43a–i) Cayratia jungii (Gregor) Chen & Manchester (Pl. 43o, p) Palaeovitis sp. (Pl. 43j–n) Unnamed Vitaceae (Pl. 43q, r) Crassivitisemen wildei (Chen & Manchester) comb. n. (Pl. 44a–m, 45a–d) MYRTALES Family: Lythraceae cf. Decodon (Pl. 25a–f, 26a–g) FABIDS [EUROSIDS I] FABALES Family: Leguminosae Mimosites spiegeli Engelhardt (Pl. 23a–f) Leguminocarpon herendeenii n. sp. (Pl. 24a–d) Leguminocarpon sp. 1 (Pl. 24e, f) Leguminocarpon sp. 2 (Pl. 24g) FAGALES Family: Juglandaceae Cruciptera schaarschmidtii Manchester, Collinson & Goth (Pl. 20d–g, 21b) Hooleya sp. (Pl. 21a) Palaeocarya sp. sensu Manchester et al. (Pl. 20h, i) MALPIGHIALES Family: Euphorbiaceae Euphorbiotheca gothii sp. n. (Pl. 10a–l, 11a–p) OXALIDALES Family: Elaeocarpaceae Sloanea messelensis sp. n. (Pl. 9a–o). ROSALES (sensu APGIII, including Urticales, Rhamnales) Family: Rhamnaceae Berchemia mellerae sp. n. (Pl. 36h–n) Family: Ulmaceae Cedrelospermum leptospermum (Ettingshausen) Manchester emend. Wilde & Manchester (Pl. 41a–f) Family: Cannabaceae Aphananthe cf. tenuicostata Dorofeev (Pl. 7i–l) MALVIDS [EUROSID II] Family: Tapisciaceae Tapiscia pusilla (Reid & Chandler) Mai (Pl. 39a–l) SAPINDALES Family: Anacardiaceae Anacardium germanicum Manchester, Wilde & Collinson (Pl. 2a–e) Lannea hessenensis sp. n. (Pl. 2g–i) Pentoperculum minimus (Reid & Chandler) Manchester (Pl. 3a–h)

Pleiogynium mitchellii sp. n. (Pl. 3i–m) Family: Burseraceae Canarium sp. (Pl. 7a–f) Bursericarpum sp. (Pl. 7g, h) Family: Rutaceae Rutaspermum chandleri Collinson & Gregor (Pl. 37m) Rutaspermum messelense Collinson & Gregor (Pl. 37a–e) Rutaceae undetermined A sensu Collinson & Gregor (Pl. 37f) Rutaceae undetermined B sensu Collinson & Gregor (Pl. 37g–i) cf. Toddalia sp. (Pl. 37j–l) Family: Simaroubaceae Ailanthus confucii Unger (Pl. 38l, m) ASTERIDS CORNALES Family: Alangiaceae Alangium sp. (Pl. 1d) Family: Mastixiaceae cf Beckettia Reid & Chandler (Pl. 29h–l) Eomastixia cf. rugosa Chandler emend. Chandler (Pl. 28a–m 29a–g) ?Mastixia sp. (Pl. 27h–t) Mastixiopsis nyssoides Kirchheimer (Pl. 30a–i) Indet. mastixioid. (Pl.s 29 n, o, 30j, k) Family: Nyssaceae Nyssa disseminata (R. Ludw.) Kirchheimer (Pl. 36a–d) ERICALES Family: Theaceae Camelliacarpoidea messelensis sp. n. (Pl. 40a–i) Family: Pentaphylacaceae Cleyera sp. (Pl. 36e–g)


Family: Sapotaceae cf. Sapotispermum sp. (Pl. 38j, k) EUASTERIDS [LAMIIDS] Family: Icacinaceae Palaeohosiea bilinica (E ttingshausen ) K vaček & Bůžek (Pl. 15a–k, m, n, 16a–c) Palaeohosiea sp. (Pl. 16d–j) cf. Natsiatum sp. (Pl. 16k–m) Palaeophytocrene sp. (Pl. 17j–p) Icacinicarya densipunctata sp. n. (Pl. 19f–k) Icacinicarya tiffneyi sp. n. (Pl. 17a–i) Icacinicarya sp. (Pl. 20a–c) Phytocrene punctilinearis sp. n. (Pl. 18a, b, d, e, h–j) Pyrenacantha sp. 1 (Pl. 18c, f, g). Pyrenacantha sp. 2 (Pl. 18k, l; 19a–d) cf. Pyrenacantha (Pl. 19e) GENTIANALES Family: Apocynaceae Cypselites sp. (Pl. 4a–e) LAMIALES Family: Bignoniaceae Darmstadtia biseriata gen. et sp. n. (Pl. 6a–c, e) CAMPANULIDS [EUASTERIDS II] Family: Toricelliaceae Toricellia bonesii (Manchester) Manchester (Pl. 39m–p) INCERTAE SEDIS Carpolithus callosaeoides (Engelhardt) comb. n. (Pl. 45e–l) Saportaspermum sp. (Pl. 46a–h) Spirellea sp. (Pl. 46i–m) Carpolithus 62 spp. (Pl. 47–76)

Modified somewhat from strict APG III circumscriptions because we retain Mastixiaceae Van Tiegh. and Nyssaceae Dum. as distinct families from Cornaceae. 1

87

Collinson, M., Manchester, S. & Wilde, V.: Fossil Fruits and Seeds of the Middle Eocene Messel biota, Germany Table 2: Representation of the different plant organs in the Messel flora. Family

Leaves

Fruits/seeds

Alangiaceae

1

Altingiaceae

1

Anacardiaceae

4

Apocynaceae

>4

Pollen

Flowers

4 7

3

Araliaceae

2

Arecaceae

1

Berberidaceae

1

3

2 1

2

2 2

2

2

1

Bombacaceae

1 3

Burseraceae

3

2

2

Buxaceae

1 (wood)

Cannabiaceae

1

1

1

Cyrillaceae

1 2

2

2

2

1 1

Eucommiaceae Euphorbiaceae

1 2

2

1

1

1

Fagaceae Hamamelidaceae

1 5 or more

5

3

2-?3

3

11

2

11

13 or more

Icacinaceae

?1

Illiciaceae

?1

Juglandaceae

4 or more

3

Lauraceae

21 or more

3 or more

1

?1

Leguminosae

>5

Loranthaceae

2

4 1

Magnoliaceae

2 or more

2 or 3

3 2 3

6

Menispermaceae

1

Moraceae

?1

Myricaceae

2

Myristicaceae

1 5

3

Mastixiaceae

13

2

Lythraceae

Myrtaceae

1

21

Lecythidaceae

Malvaceae

1

1

Elaeocarpaceae

1 1

1

Cyclanthaceae

Ericaceae

1

1

Chloranthaceae Cyperaceae

2 1

Betulaceae Bignoniaceae

minimum richness 1 1

?1

1

Aquifoliaceae

Cercidiphyllaceae

Other

2

6

17

17 1 2

2

1

1

1

1

Nyctaginaceae

?1

?1

1

Nymphaeales

2

?1

1

?1

2

Nyssaceae

1

1

2

2

Olacaceae

2

2

Oleaceae

1 or 2

2

Platanaceae

1

1

Restionaceae

2

Rhamnaceae Rosaceae

88

1 1

2 1

?1

1


Family

Leaves

Fruits/seeds

Pollen

Flowers

Rutaceae

1

5

?4

1

Sabiaceae

1 or 2

Salicaceae Sapindaceae Schisandraceae

1

1

1

Tapisciaceae

1 ?1

1

1

1

1 5

1

1

5

Thymelaeaceae

1

Tiliaceae

2

Toricelliaceae Ulmaceae

1 2

1

Verbenaceae Vitaceae

1

1-2 1

1

Sterculiaceae

Theaceae

1

1

Simaroubaceae Symplocaceae

1 ?1

1

minimum richness 5 2

1

Sapotaceae

Other

7

1 1

2 1

4

4

?1

1

2

7

Based on Wilde (2004) and the present monograph. Main sources as follows: Leaves – Wilde (1989), Wilde et al. (2005, Araceae), Kvaček (1988, Lauraceae), Kvaček & Wilde (2010, Malvales), Wilde & Frankenhäuser (2000, Myricaceae), Wilde & Manchester (2003, Ulmaceae); Fruits and seeds – herein; Pollen – Thiele-Pfeiffer (1988); Flowers – Schaarschmidt (1984, 1986), Wilde & Schaarschmidt (1993), Schaarschmidt & Wilde 1986 and Harley 1997 (Arecaceae); Wood – Wilde & Süss (2001).

89

Collinson, M., Manchester, S. & Wilde, V.: Fossil Fruits and Seeds of the Middle Eocene Messel biota, Germany Table 3: Tissue and organ preservation in the angiosperm fruit and seed flora from Messel noting features relevant to dispersal biology and having significance for animal diet*.

Taxon

Exceptional organic preservation

Significance for seed dispersal or for animal feeding

Comments, type of specimen

Family Alangiaceae

Alangium sp.

fruit w endocarp

Family Altingiaceae

Steinhauera subglobosa

infructescence, fruit wall complete

dry seeds in capsule

Family Anacardiaceae

Anacardium germanicum

fruit wall complete; hypocarp

fleshy hypocarp

germination valve

Lannea hessenensis

fruit wall complete

Pentoperculum minimus

fruit wall complete

Pleiogynium mitchellii

fruit wall complete

Family Apocynaceae

Cypselites sp.

hair tuft (plume, coma)

wind dispersal/seed orientation

Family Arecaceae

Friedemannia messelensis

fruit complete. persistent calyx

thin outer fruit wall + endocarp

fibrous fruit wall tissue

Family Bignoniaceae

Darmstadtia biseriata

dry dehiscent fruit

unwinged seeds or wing not preserved

Family Burseraceae

Canarium sp.

Bursericarpum sp.

Family Cannabaceae

Aphananthe cf. tenuicostata

endocarp only

Family Cyclanthaceae

Cyclanthus messelensis

fruiting cycle

tiny dry seeds, 2 mm long

Family Cyperaceae

Volkeria messelense

infructescence

small dry fruits

attached stamens w pollen

Family Elaeocarpaceae

Sloanea messelensis

fruit wall complete

outer fruit wall thin + endocarp

Family Euphorbiaceae

Euphorbiotheca gothii

fruit wall complete

dry dehiscent, likely explosive

dry seeds released

Family Hamamelidaceae

Mytilaria boglei

leafy shoots attached infructescences

capsules

immature

Corylopsis maii

infructescence

capsules, explosive, dry seed released

Corylopsis waltheri

dry seeds, possibly explosive release

Family Icacinaceae

90

thin outer fruit wall + endocarp, fibrous thin outer fruit wall + endocarp thin outer fruit wall + endocarp

germination valve germination valve germination valve

endocarp only, germination valve endocarp only, germination valve


Taxon




Palaeohosiea bilinica

endocarp only

cf. Natsiatum sp.

endocarp only

Palaeohosiea sp.

endocarp only

Palaeophytocrene sp.

endocarp only

Phytocrene puctilinearis

endocarp only

Pyrenacantha sp. 1

outer fruit wall

thin fleshy tissue + endocarp

Pyrenacantha sp. 2

endocarp only

cf. Pyrenacantha

endocarp only endocarp only

Icacinicarya densipunctata Icacinicarya tiffneyi

outer fruit wall

thin fleshy tissue + endocarp

thin, unclear outer tissues

Icacinicarya sp.

endocarp only

Family Juglandaceae

Cruciptera schaarschmidtii

helicopter wing

wind dispersal

Hooleya sp.

two lateral wings

wind dispersal

Palaeocarya sp.

trilobed wing, small 4th lobe

wind dispersal

Family Lauraceae

Laurocarpum sp. 1

outer fruit wall

possible leathery softer tissues

Laurocarpum sp. 2

outer fruit wall

possible leathery softer tissues

Laurocarpum sp. 3

outer fruit wall

thick (3 mm) mesocarp

Lauraceae genus indet.

outer fruit wall

no detail observed

Family Leguminosae

Mimosites spiegeli

complete pod

dry pod, dehiscence not proven

inferred dry seeds

Leguminocarpon herendeenii

complete pod

dry pod, dehiscence suggested

inferred dry seeds

Leguminocarpon sp. 1

complete pod

Leguminocarpon sp. 2

complete pod

Family Lythraceae

cf. Decodon

infructescence

dry seeds

Family Magnoliaceae

Magnolia sp. 1

multifollicle

Magnolia spp.

dry seeds

Family Mastixiaceae

cf. Beckettia

outer fruit wall

thin mesocarp + endocarp

germination valve

Eomastixia cf. rugosa

fibrous tissue, resin canals

endocarp, valve not obvious

?Mastixia sp.

possible outer fruit wall

thin mesocarp + endocarp

might be in coprolite, germination valve

Mastixiopsis nyssoides

woody and fibrous tissue

endocarp germination valve

Indet. mastixioid

patches of epicarp

Thin mesocarp + endocarp

germination valve

Family Menispermaceae

Diploclisia rugulosa

endocarp only

dry pod, dehiscence not proven dry pod, dehiscence not proven

inferred dry seeds inferred dry seeds

91


Taxon




Stephania hootae

endocarp only

Indeterminate Menispermeae

endocarp only

Martinmuellera tuberculata

some clusters of endocarps

coprolite and fruiting panicle

Palaeosinomenium ornamentum

endocarp only

Palaeosinomenium venablesii

endocarp only

Palaeosinomenium sp.

endocarp only

?Pericampylus sp.

endocarp only

Wardensheppeya sp.

endocarp only

Karinschmidtia rotulae

fruit epicarp; wing like extension

thick (4 mm) soft mesocarp + endocarp

Cocculus lottii

endocarp only

Tinomiscoidea jacquesii

endocarp only

Tinosporeae sp. 1

endocarp only

Parabaena cf. europaea

endocarp only

Tinosporeae sp. 2

endocarp only

Tinosporeae sp. 3

endocarp only

Unnamed Menispermaceae

endocarp only

Myristicacarpum sp.

ruminate endosperm

endosperm well-developed

seed

Order ? Nymphaeales

Unnamed species.

mature flowers with seeds

soft fruit tissues

Detailed study in progress

Family Nyssaceae

Nyssa disseminata

endocarp only, germination valve

Family Pentaphylacaceae

Cleyera sp

small seed

Family Rhamnaceae

Berchemia mellerae

portion of outer fruit wall

Soft tissue c. 1.5 mm thick

germination area

Family Rutaceae

Rutaspermum chandleri

seed

Rutaspermum messelense

seed

Rutaceae undetermined A

seed

Rutaceae undetermined B

seed

cf. Toddalia sp.

seed

Family Sabiaceae

Meliosma sp.

endocarp only

Family Sapotaceae

cf. Sapotispermum sp.

large seed, thick seed coat

Family Myristicaceae

92


Taxon




Family Simaroubaceae

Ailanthus confucii

two lateral tapering wings

wind dispersal

Family: Tapisciaceae

Tapiscia pusilla

outer fruit wall; clusters

thin outer layer + seed;

cluster are probably coprolites

Family Theaceae

Camelliacarpoidea messelensis

fruit complete, persistent calyx

dry capsule, dry brittle seeds released

Family Toricelliaceae

Toricellia bonesii

endocarp only, germination valve

Family Ulmaceae

Cedrelospermum leptospermum

samara, leafy shoots attached fruits

wind dispersal

Family Vitaceae

Vitis messelensis

partial fruits with seeds

soft tissues

seeds, some coprolites

Parthenocissus britannica

partial fruits with seeds

soft tissue c. 1 mm thick

seeds

Ampelopsis sp.

seed

Cayratia jungii

seed

Palaeovitis sp.

seed

Unnamed Vitaceae

seed

Crassivitisemen wildei

complete fruits

soft tissue up to 3 mm thick

large single seed

Carpolithus callosaeoides

thick woody tissues

fruit

Saportaspermum sp.

winged seed

wind dispersal

Incertae Sedis

small seed

Carpolithus sp. 1

dry valvate

fruit

Carpolithus sp 2

awned fruit

Carpolithus sp 3

Carpolithus sp 4

aid dispersal/anchorage fibrous tissue, softer not woody

Carpolithus sp 5

dry, winged seeds

berry soft fruit and seed tissues

Carpolithus sp 7

large multiseeded fruit epicarp, crumpled fruit and seed

dry capsule

Carpolithus sp 8

cluster

seed/nutlet

Carpolithus sp 9

small fruit

Carpolithus sp 10

possible berry

fruit seeds protruding

Carpolithus sp 11

uncertain

Carpolithus sp 12

dry seeds

fruit

Carpolithus sp 13

leathery capsule

many seeded berry

Carpolithus sp 15

seed cluster with organic surround large fruit 44 mm

large leathery tissues

Carpolithus sp 16

infructescence

dry

Carpolithus sp 17

infructesence fruit crumpled

possible softer tissues

Carpolithus sp 18

very large fruit 64x46 mm

leathery to fleshy, brittle seeds

large seeds 19x16 mm

Carpolithus sp 19

thin membraneous fruit

likely soft fleshy tissues

Carpolithus sp 20

fin winged fruit

wind dispersal

Spirellea sp.

Carpolithus sp 6

Carpolithus sp 14

fruit seed

93


Taxon



Carpolithus sp 21

cluster tiny winged seeds

wind dispersal

Carpolithus sp 22

protection or anchorage

Comments, type of specimen seed body 1.5 mm, wing 1.8 mm spiny fruit

Carpolithus sp 23

dehiscent dry

fruit

endosperm well developed

seed

fruit

wind dispersal

seed body 1.3 mm

Carpolithus sp 27

possible ruminate endosperm seed w very fine long spreading hairs racemose infructescence

dry fruits

Carpolithus sp 28

seed

Carpolithus sp 29

leathery valves

berry with soft tissues

Carpolithus sp 31

infructescence multiseeded fruit large 33 mm

seed

Carpolithus sp 32

?leathery

fruit

Carpolithus sp 33

infructescence

Carpolithus sp 34

possible soft tissues

Carpolithus sp 35

cluster

Possible fruit tissues

immature maybe coprolite, possible crystal cells seed/endocarp

Carpolithus sp 36

globose infructescence white viscous spongy contents hairy fruit with resin membraneous valves or wings infructescence

woody, dry capsule

thick (3 mm) woody wall

fruit

possible protection

unusual characters

possible dispersal

seeds usually found isolated

berry-like

fruit

uncertain special function?

fruit

Carpolithus sp 43

unusual pericarp tissue layer very large fruit 92x60 mm

brittle surround to softer tissue

Carpolithus sp 44

uncertain

Carpolithus sp 45

valvate dry

Carpolithus sp 46

softer tissues

softer tissues large 34 mm

Carpolithus sp 47

fruit crumpled cuticle & inner tissue ?seed

Carpolithus sp 48

small dry

fruit

Carpolithus sp 49

infructescence

valvate dry

Carpolithus sp 50

seed

Carpolithus sp 51

seed

Carpolithus sp 52

?fruit

very soft tissues

softest tissues at Messel

Carpolithus sp 54

pericarp cuticle, many small seeds

Large Woody

Large fruit 28x36 mm

Carpolithus sp 55

circular wing

wind dispersal

four fruits on wing

Carpolithus sp 56

one cluster

berry, tiny seeds

might be a coprolite

Carpolithus sp 57

In Masillamys gut content

small brittle shiny seeds

Carpolithus sp 58

aril

Possible insect dispersal

seed

Carpolithus sp 59

fruit with hypanthium

Carpolithus sp 60

Carpolithus sp 61

long folded & crumpled wings

valves, ?leathery leathery outer, softer inner tissue dry

Carpolithus sp 24 Carpolithus sp 25 Carpolithus sp 26

Carpolithus sp 30

Carpolithus sp 37 Carpolithus sp 38 Carpolithus sp 39 Carpolithus sp 40 Carpolithus sp 41 Carpolithus sp 42

Carpolithus sp 53

Carpolithus sp 62

wind dispersal

fruit 25x17 mm fruit fruit with basally attached wings

* The term woody is here used for strongly sclerified tissue, such as in the endocarp of modern Juglans; the term fibrous when fibres are observed in the specimens and the term leathery for textures that are judged comparable to those of the modern Aesculus fruit wall or where thick-walled tissue in Messel specimens is crumpled or wrinkled.

94


Plates 1–76

95


Plate 1 Doliostrobaceae, Alangiaceae a–c: Doliostrobus taxiformis (Sternberg) Z. Kvaček emend. Kunzmann, a: SM.B Me 20102, cone scale with straight lateral margins and protruding apical tip; b: SM.B Me 20101, cone scale with concave lateral margins; c: SM.B Me 20100, narrower scale; d: Alangium sp., SM.B Me 2215, fruit with two carpels separated by a longitudinal groove accommodating a row of circular depressions; e–f: Steinhauera subglobosa Presl, emend. Mai, SM.B Me 19093, e: at least seven subglobose flowering/fruiting heads borne along a common axis; f: detail of the largest fruiting head in showing bilocular capsules, lacking persistent styles, embedded in smooth tissue; scale bars: a–c, f = 1 cm; d = 5 mm; e = 2 cm.

96


Plate 2 Anacardiaceae a–e: Anacardium germanicum Manchester, Wilde & Collinson, a: SM.B Me 7156, fruit with enlarged pedicel (hypocarp); b: holotype, SM.B Me 7139, showing curved fruit and persistent hypocarp; c: SM.B Me 17770, fruit with a relatively elongate hypocarp; d: SM.B Me 18095, specimen with some layers of fruit wall fractured away; e: detail from d of exocarp and underlying mesocarp; f: Extant fruit of A. occidentale, fruit and hypocarp in lateral surface view, Clarendon, Jamaica. BM Herbarium, Coll. Hope 7633; g–i: Lannea hessenensis sp. n., g: SM.B Me 5583, outer endocarp surface covered with fine swirling fibres and oblique apical slit marking the germination valve (arrows); h: holotype, SM.B Me 4882, endocarp with fibrous strands enclosing large deep depressions, with oblique apical germination valve (arrows) and basal attachment scars; i: SM.B Me 15856, endocarp with sculpture including smooth, sunken areoles. Note oblique apical germination valve; j: Extant Lannea coromandelica (Houtt.) Merr. endocarp with germination valve (arrows). US Herbarium, 258427A, Coll. C. J. Saldenha 16725, Hassan District, India; scale bars: a–d, f–j = 1 cm; e = 2 mm.

98


Plate 3 Anacardiaceae a–h: Pentoperculum minimus (Reid & Chandler) Manchester, a: SM.B Me 4771, oblique apical view of specimen showing pentagonal outline and longitudinal lines of germination valve splitting; b: SM.B Me 21467, viewed basally showing pentagonal outline with five major ribs with intervening minor ribs; c: M.B Me 21279, lateral compaction showing the apical plugs (arrows) and two of the paired depressions; d: SM.B Me 11234, lateral compaction showing two of the 10 elliptical depressions; e: SM.B Me 2098, transversely broken specimen clearly shows the radially arranged locules alternating with thick lacunate septa; f–h: SM.B Me 8810, apical, basal and lateral view, respectively, probably foreshortened through compaction; note bulging elliptical germination valves radiating from center of apical side; i–m: Pleiogynium mitchellii sp. n., holotype, SM.B Me 2227, i: apical view, showing circular outline, and 10 radiating ridges, each with a bulging elliptical germination valve; j: basal view with central depression of pedicel scar; k: equatorial view, showing broadly rounded apical side and widely conical basal side; l: digital SRXTM vertical section, showing two elongate locules; m: digital SRXTM transverse section, showing radially arranged locules and vascular bundles; n, o: Extant Pleiogynium timoriensis (DC) Leenh. GH Herbarium, coll. A. C. Smith 1499 from Fiji, side and apical views; p: P. timoriensis NY Herbarium, coll. P. A. Cox 1113 from Tonga, transverse section of the fruit for comparison with m; scale bars: a–h = 5 mm; i–p = 1 cm.

100


Plate 4 Apocynaceae a–e: Cypselites sp., a: SM.B Me 7624, seed with terminal elongated tuft of hairs; b: enlargement of a, showing seed body longitudinally striated and rectangular cells in longitudinal rows; c: SM.B Me 20180, seed body with pointed base; d: SM.B Me 16422, large example; e: SM.B Me 15469, seed body longitudinally striated and with pointed base; scale bars: a–d = 1 cm; e = 5 mm.

102


Plate 5 Arecaceae a–n: Friedemannia messelensis gen. et sp. n., a: SM.B Me 2380, fruit with persistent calyx and style; b: SM.B Me 2134, partially detached calyx; c: SM.B Me 2130, fruit with persistent calyx; d: holotype, SM.B Me 2135, broad complete fruit with persistent calyx, style and fibrous outer wall; e: SM.B Me 2145, narrow fruit with well preserved persistent calyx lobes; f: SM.B Me 2123, fruit wall fibres spreading as a result of degradation; g: SM.B Me 2378, fruit with fibrous persistent calyx and apical scars (? fungal/insect damage); h, i: SM.B Me 2131, fruit with persistent calyx showing short and long lobes; j–l: fruits revealing internal structure; j: SM.B Me 2137, fruit with persistent calyx, outer fibrous wall and terminal style; fruit cut with a razor revealing terminal ‘scar’ of hilar chalazal plug and fibrous bands radiating around raphe; k, l: SM.B Me 2148, SM.B Me 2147, natural breaks in specimens revealing terminal hilar chalazal plug and fibre bands radiating around raphe; m: SM.B Me 2138, seed showing the hilar/chalazal plug through the partially degraded fruit wall; n: SM.B Me 2114, young fruit showing two calyx whorls; scale bars: a–n = 1 cm.

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Plate 6 Bignoniaceae a–c, e: Darmstadtia biseriata gen. et sp. n., a, c: SM.B Me 7140, fruit with portions of the fruit wall detached, revealing central placental strand and elliptical seeds within; b, e: holotype, SM.B Me 7143, fruit with portion of facing valve shed, revealing two rows of seeds; d, f: Extant fruit of Ceratophytum tetragonolobum (Jacq.) Sprague & Sandw., for comparison GH Herbarium, coll. Gentry & Morillo 10349; scale bars: a–f = 1 cm.

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Plate 7 Burseraceae, Cannabiaceae a–f: Canarium sp., a–d: SM.B Me 2300; a: elliptical endocarp in lateral view showing two flat faces joining in a longitudinal ridge; b: specimen turned over showing the third face; c, d: apical and basal views of a, b, showing trigonal outline and narrowly grooved keels; e: HLMD-Me-15716, oblique lateral view showing truncate base, rounded lateral margins, and a longtitudinal ridge down length of where two lateral faces meet; f: SM.B Me 4610, endocarp from which the apical triangular germination valve has been shed revealing smooth locule with a pointed apex; g, h: Bursericarpum sp., g: SM.B Me 23208, endocarp in lateral view with smoothly curved dorsal margin (left) and base, and prominent placental slit on the ventral side; h: SM.B Me 18255, D-shaped endocarp in lateral view, with straight ventral margin (left), curved dorsal margin (right) and transverse placental slit (arrow); i–l: Aphananthe cf. tenuicostata Dorofeev, i: SM.B Me 4169, endocarp with longitudinal fold suggesting originally pyriform shape, subapical protrusion uncrushed; j: SM.B Me 4466, ovate endocarp with prominent, asymmetrically placed subapical protrusion; wall lined with papery shiny cuticle; k: SM.B. Me 13049, endocarp in lateral view with subapical protrusion; l: SM.B Me 4476, specimen removed from oil shale clearly showing subapical protrusion; scale bars: a–f = 1 cm; g, h = 5 mm; I–k = 4 mm; l = 3 mm.

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Plate 8 Cyclanthaceae, Cyperaceae a–e: Cyclanthus messelensis S. Y. Smith, Collinson & Rudall, a: SM.B Me 2005, discoidal fruiting cycle with a central hole, radiating fibre strands, and a thickened outer rim; b: SM.B Me 24734, incomplete discoidal fruiting cycle showing elliptical seeds (arrows) between cuticle layers; c: detail from a, showing fibre strands and included seeds (arrows); d, e: Same, as b, enlarged, showing details of longitudinally ribbed seeds; f–i: Volkeria messelense Smith, Collinson, Simpson & Rudall, f: SM.B Me 2107, pedunculate infructescence; g: SM.B Me 2104, infructescence viewed from base; h: SM.B Me 4313, isolated fruit, surface view; i: SM.B Me 4317, isolated fruit fractured showing locule and surrounding thick wall; scale bars: a, b = 1 cm; c, d, f = 5 mm; e = 3 mm; g–i = 1 mm.

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Plate 9 Elaeocarpaceae a–o: Sloanea messelensis sp. n., a: SM.B Me 7149, dehisced, paired valves showing smooth locule surface and surrounding spiny wall; b: SM.B Me 23881, fruit with complete outer wall compressed so as to reveal the circular basal disk (arrow); c: SM.B Me 7082, fruit with a long pedicel in lateral view; d: holotype, SM.B Me 7133, fractured fruit with a subtending disk (arrow), and a long pedicel; e: SM.B Me 18161, pedicellate mature fruit with intact surface; f: SM.B Me 20053, pedicellate immature fruit with prominent hypogynous perianth; g: SM.B Me 20449, developing fruit retaining terminal style and prominent hypogynous perianth; h: SM.B Me 20039, fruit portion showing smooth ventral surface and surrounding spiny wall; i: SM.B Me 2052, isolated valve, viewed ventrally, with median ridge representing the septum; j: SM.B Me 17898, partially broken fruit with excellent preservation showing very fine closely spaced conical spinules; k: SM.B Me 7163; fruit showing basal disk and meridional grooves separating the valves; l: top view from k of obliquely fractured fruit showing compacted fruit with relatively thin pericarp; m: SM.B Me 17775, oblique-basal view showing circular outline of the fruit and basal scar; n: SM.B Me 7179, oblique-apical view, showing radially oriented bulges at the periphery that may indicate five carpels; o: SM.B Me 16420, portion of fruit showing the columnar arrangement of tissues leading to the peripheral spines, and smooth locules; scale bars: a–o = 1 cm.

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Plate 10 Euphorbiaceae a–l: Euphorbiotheca gothii sp. n., a: SM.B Me 16561, complete fruit with six segments on a stout pedicel attached to a twig, probably immature; b: SM.B Me 4718, fruit transversely compressed, radially symmetrical with meridional grooves delimiting six valves; c: SM.B Me 17492, obliquely compressed, partially dehiscent fruit showing six valves, with leathery exterior, and smooth interior valve margins. The segments are in three pairs, revealing plane of loculicidal followed by septicidal dehiscence; d–l: various appearances resulting from dehiscence revealing the endocarp and partial separation into cocci; d: holotype, SM.B Me 7117, a pair of valves separating loculicidally, showing smooth band of endocarp tissue surrounding the locular area; e: SM.B Me 7362, paired segments, with mesocarp split apart, but still joined by endocarp; f: SM.B Me 19537, pair of dehisced segments with apical parts of endocarps torn away from the mesocarp; g: SM.B Me 21681, dehiscent fruit with apical protrusions of the endocarp (arrows) separating from the mesocarp; h: SM.B Me 21677, transversely compressed fruit, with three paired segments, indicating loculicidal splitting in progress, prior to septicidal splitting; i: SM.B Me 21678, shed, bivalved coccus corresponding to one of the units seen in h; j: SM.B Me 8264, pair of cocci attached by the basal part of the endocarp; k: SM.B Me 8076, isolated coccus, showing well-defined endocarp lining the locule; l: SM.B Me 13838, partially dehisced fruit showing displacement and twisting of three loculicidally separated segments; scale bars: a = 1 cm; b–l = 5 mm.

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Plate 11 Euphorbiaceae a–p: Euphorbiotheca gothii sp. n., a: SM.B Me 21531, fruit viewed obliquely from the base, showing stout pedicel and six radially symmetrical segments; b: SM.B Me 7581, two adhering cocci, showing the darker endocarp with T-shaped junctions at the apex of locule (arrow); c: SM.B Me 7185, single coccus, ventral surface, with seed and endocarp missing; d: SM.B Me 7262, single coccus, dorsal surface; e: SM.B Me 17552, coccus in lateral view showing endocarp, mesocarp and exocarp; note T-shape ventralapical extension of endocarp (arrow), detail of locule lining can also be observed in this specimen; f: SM.B Me 7285, paired dehiscent segments, with endocarp missing from left valve, yet still intact on the right valve; g: SM.B Me 8063, pair of cocci, twisted with the apices facing opposite directions; h: SM.B Me 4799, isolated endocarp completely separated from mesocarp; i: SM.B Me 7268, single coccus, with convex external surface at right; j: SM.B Me 7358, single segment, with protruding remains of endocarp; k: SM.B Me 8297, portion of fruit with one valve shed, showing impressions of two seeds within a pair of locules; l: enlargement of k showing outlines of two seeds and their placentation; m: SM.B Me 17327, adjacent cocci exposed by loss of a third coccus, showing circular outline of locule, anticlinal arrangement of fibers in the mesocarp, and thickened band of endocarp tissue surrounding lining of the locule; n: SM. Me 17044, coccus with elliptical seed in situ; o: same as n, enlarged, showing elliptical seed surrounded by band of endocarp tissue; p: seed from o, enlarged showing fine cellular surface pattern; scale bars: a–o = 5 mm; p = 2 mm.

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Plate 12 Hamamelidaceae a: Mytilaria boglei sp. n., holotype, SM.B Me 18788, branch with attached leaves and fruits; details presented in pl. 13; scale bar: a = 5 cm.

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Plate 13 Hamamelidaceae a, b, d–f: Mytilaria boglei sp. n., holotype, SM.B Me 18788, a: infructescence from leaf-bearing twig of pl. 12, showing unequally developed fruits, and protrusion of distal end of raceme lacking developed fruits; b: infructescence from same twig as a, showing unequally developed fruits, and protrusion of distal end of raceme lacking developed fruits; c: SM. B Me 21325, isolated dispersed infructescence; d: enlargement of the infructescence in a with protrusion of raceme with undeveloped fruits toward top of the image; e: enlargement of raceme from d, showing helical arrangement of florets with domed gynoecia; f: detail from e, showing bipartite diamond-shaped outline and domed surface of individual gynoecia and the hypanthium tissue separating adjacent florets; g–m: Extant Mytilaria laosensis H. Lecompte for comparison, g: herbarium sheet showing arrangement of lobed leaves on twig. A Herbarium, coll. K. M Feng 11447, southeastern Yunnan, China; h: raceme showing helically arranged gynoecia. A Herbarium, coll. W. T. Tsang 24730, Kwangsi, China; i: enlarged fruit with distorted remains of raceme. A Herbarium, coll. K. M Feng 11558, southeastern Yunnan, China; j, k: lateral (j) and apical (k) view of matured, dehisced capsular fruits. A Herbarium, coll. W. T. Tsang 24098, Kwangsi, China; l: seeds from the same collection as j, k; m: detail of raceme from h, showing domed gynoecia, and surrounding hypanthium tissue for comparison with e, f; scale bars: a–d, h–m = 1 cm; e, f = 5 mm; g = 10 cm.

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Plate 14 Hamamelidaceae a–e: Corylopsis maii sp. n. infructescence, a: SM.B Me 7434, infructescence with a stout axis and numerous sessile fruits; b: SM.B Me 16263, infructescence with rounded bilaterally symmetrical sessile fruits; c: holotype, SM.B Me 14508, infructescence with cup-like structures surrounding the base of fruits (arrows) and persistent pair of styles; d: SM.B Me 14383, single loculicidally dehisced capsule showing pair of locules and apical septicidal groove. e: SM.B Me 23961, infructescence with apex of fruits with paired persistent styles with divergent tips (arrows); f–t: Corylopsis waltheri sp. n. seeds, f: SM.B Me 4722, holotype showing light-colored ligulate hilar scar on otherwise dark, somewhat shiny seed; g: SM.B Me 7129, similar seed with hilum outline faintly visible in apical half; h: SM.B Me 2177, large seed with elongate, recessed hilar scar near apex; i–k: SM.B Me 20522, single seed removed from oil shale viewed in different orientations, showing that hilar scar (j, k) reaches, but does not pass over the apical end; l, m: SM.B Me 2181; n: SM.B Me 7939; o: SM.B Me 7922; p: SM.B Me 4726; q, r: SM.B Me 7127; s: SM.B Me 7933; t: SM.B Me 5143, a long narrow hilar scar covering over the apex; scale bars: a–c, e = 1 cm; d, f–t = 5 mm.

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Plate 15 Icacinaceae a–k, m, n: Palaeohosiea bilinica (Ettingshausen) Kvaček & Bůžek, a: SM.B Me 4919, wide elliptical endocarp with typical reticulate surface pattern; cracks are unusual for this taxon; b: SM.B Me 2081, ovate endocarp, showing blind-ending ridges within areoles of the surface reticulum; c: SM.B Me 4921, endocarp with prominent funicular bulge at apex; d: SM.B Me 5625, endocarp split along the plane of bisymmetry showing relatively smooth locule and thickness of wall; e, f: SM.B ME 2091, opposite sides of single endocarp showing differences in prominence of longitudinal ridges; g: SM.B Me 7341, elliptical endocarp, with three particularly prominent longitudinal ridges and reticulate pattern with occasional blind-ending ridges; h: SM.B Me 2080, endocarp with three prominent longitudinal ridges; i: SM.B Me 17596, endocarp with obvious blind-ending branches from the ridges; j: SM.B Me 2086, broadly elliptical endocarp; k: SM.B Me 4616, damaged specimen with intact outer reticulum exposed in the lower half, smooth locule and broken endocarp wall seen in upper half; l: Endocarp of extant Natsiatum herpeticum (A: Grierson & Long 3775, Bhutan); m, n: Palaeohosiea bilinica, SM.B Me 2357, unilocular endocarp, locule wall smooth (papillae lost on exposed areas), SEM; scale bars: a–l = 5 mm; m, n = 2 mm.

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Plate 16 Icacinaceae a–c: Palaeohosiea bilinica (Ettingshausen) Kvaček & Bůžek, SM.B Me 4652, portion of an obliquely fractured endocarp, a: reticulate endocarp surface and thick wall; b: detail of broken locule surface showing overlapping tissue layers; c: detail from b, showing a layer of papillae (left) overlying the layer of smaller isodiametric cells (lower right); d–j: Palaeohosiea sp., d: SM.B Me 4089, endocarp with obtusely pointed apex; e: SM.B Me 4626, broad endocarp with longitudinally elongate reticulate fields towards the apex; f: SM.B Me 21587, endocarp with rounded base; note grooves at the ridge crests; g: SM.B Me 17702, endocarp with ovate outline; h: Detail from small fragment of e, broken open to expose inner wall thickness and locule lining; i: fragment removed from center of h, showing widely spaced, circular papillae lining the locule, SEM; j: vertical section of fruit wall from fragment in i, showing pitted thick-walled sclereids, SEM; k–m: cf. Natsiatum sp., k, m: SM.B Me 4926; k: ovate endocarp in face view with reticulate surface including fine branches within the areoles; l: SM.B Me 17529, elliptical endocarp with similar surface sculpture but more rounded base and apex; m: enlargement of k, showing areoles containing numerous fine blindly ending branches from the ridges; scale bars: a = 5 mm; b, h = 1 mm; c = 0.5 mm; d–g, k, l = 1 cm; i = 100 µm; j = 30 µm; m = 4 mm.

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Plate 17 Icacinaceae a–i: Icacinicarya tiffneyi sp. n., a: SM.B Me 16650, large fruit with irregular longitudinal ribbing, and prominent, horizontally elongate subapical scar (arrows) b: SM.B Me 24055, another specimen, with apical slit (arrow); c: SM.B Me 12911, specimen showing longitudinal ribs arising from base and asymmetrical apical bulge; d: SM.B Me 5557, narrower obovate specimen; e–g: holotype, SM.B Me 5619, single specimen viewed on both sides and in transverse section showing lenticular outline and single locule filled with dark substance; h: detail of the wall from g; i: detail of fibrous of ridges forming reticulum on the endocarp surface from c; j–p: Palaeophytocrene sp., SM.B Me 21413, j: surface view of relatively small rounded elliptical endocarp with inconspicuous small shallow pits; k: detail of j showing dark brown shiny outer layer in upper part of image and around edges of surface pits into which it descends, surface pits marked by pale patches; l–p: digital SRXTM sections of j; l: median longitudinal section showing projections into locule; m: transverse section of j; locule colapsed near margins due to compaction; n: near-marginal oblique tangential section of j; o: combined biplanar digital sections of j, showing the wall in transverse (upper 1/3) and tangential (lower 2/3) sections p: detail of o showing wall thickness and hollow projection into locule (cf. pl. 19p); scale bars: a–g = 1cm; h = 4 mm; i, l–o = 2 mm; j = 5 mm; k, p = 1 mm.

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Plate 18 Icacinaceae – Phytocreneae a, b: Phytocrene punctilinearis sp. n., a: SM.B Me 17899, large endocarp showing pits arranged in longitudinal rows; b: holotype, SM.B Me 2326, large endocarp partially broken revealing thickness of the endocarp wall; c: Pyrenacantha sp. 1, SM.B Me 16691, (inset) same magnification as b for comparison; d, e: Phytocrene punctilinearis sp. n., d: SM.B Me 12279, small elliptical endocarp showing pits arranged in longitudinal rows; e: SM.B Me 7150, small endocarp showing pointed apex; f, g: Pyrenacantha sp. 1, f: SM.B Me 4067, small endocarp showing rounded base, pointed apex, pitted surface, with a pattern of even, radial, spacing of pits at the periphery; g: SM.B Me 14490, similar specimen, but with surrounding soft tissue preserved; h–j: Phytocrene punctilinearis sp. n.; SM.B Me 8449, endocarp with enlargement of broken edge showing relatively thin wall (arrows in j); k, l: Pyrenacantha sp. 2, k: SM.B Me 8464, inflated, widely elliptical endocarp with prominent evenly spaced surface pits; l: SM.B Me 21431, smaller endocarp showing features as in k; scale bars: a–h, k, l = 1 cm; i =5 mm; j = 2 mm.

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Plate 19 Icacinaceae – Phytocreneae a–d: Pyrenacantha sp. 2, SM.B Me 4992, a: widely elliptical endocarp, face view showing pits; b: lenticular in lateral view showing relatively little compaction; c, d: successive enlargements showing details of pits funneling to small vertical canals. Surface layer with isodiametric cells not descending into the pits; e: Pyrenacantha sp. 3, elliptical endocarp, surface with numerous densely and evenly spaced small pits, SM.B Me 7165; f–k: Icacinicarya densipunctata sp. n., f–h: holotype, SM.B Me 23388; f: nearly circular outline of endocarp in face view with funicular protrusion; g: lenticular in apical view, with funicular depression and protrusion; h: enlargement of f showing surface covered densely with fine pits; i–k: SM.B Me 14938; i, j: wide elliptical endocarp, front and back face views, funicular protrusion not obvious but organisation otherwise identical to f; k: enlargement of i, showing surface covered densely with fine pits; l–o: fruits of extant Icacinaceae for comparison, l, m: Pyrenacantha malvifolia [Native Tanganyika] UC Santa Barbara Greenhouse, courtesy B. H. Tiffney. External view, and cut open showing elongate narrow protrusions into the locule; n, o: Phytocrene oblonga, surface of endocarp, and cut open showing pits in the wall but lack of protrusions into the locule. A. L. Pierre 2837; p: silicified Paleophytocrene pseudopersica Scott encdocarp from the Eocene Clarno Nut Beds, cut transversely showing hollow projections into the locule (cf. pl. 17p). UF 225-8597; scale bars: a, b, e–k, n–p = 1 cm; c, l, m = 5 mm; d = 1 mm

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Plate 20 Icacinaceae, Juglandaceae a–c: Icacinicarya sp., SM.B Me 10564, a: endocarp circular in face view, originally almost globose; b: enlargment of a, showing surface ornamented with flat-topped conical protruding tubercles, each having a central crater-like pit; c: detail of b, showing fine cellular pattern covering the surface but not descending into the central pit (circular area) of each tubercle; d–g: Cruciptera schaarschmidtii Manchester, Collinson & Goth, d: holotype, SM.B Me 7152, fruit with globose central nutlet and four free strap-like wings; e: SM.B Me 17520, fruit with smaller nut and symmetrical wings; f, g: SM.B Me 20099, fruit with relatively wide wings; g: enlargement of f, showing subparallel wing venation and lack of thickened midvein; h, i: Palaeocarya sp. sensu Manchester et al. 1994, h: SM.B Me 8912, pedicellate nut with trilobate bract-wing; i: SM.B Me 8694, pedicellate nut showing thickened midvein in wing and two of the three lobes of the main wing, plus shorter prophyllum enclosing the nut; scale bars: a, d–f, h, i = 1 cm; b = 1 mm; c = 0.5 mm; g = 2 mm.

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Plate 21 Juglandaceae, Lauraceae a: Hooleya sp., SM.B Me 2064, showing nut with pair of lateral wings; b: cluster of Cruciptera schaarschmidtii winged fruits, SM.B Me 24511; c, d: Lauraceae genus indet., SM.B Me 8375, c: small globose fruit with an abrupt acute apex; d: enlargement of c, showing dark smooth cuticle through which can be seen numerous yellow dots (interpreted to represent oil cell contents); e–g: Laurocarpum sp. 3, SM.B Me 18005, e: broken specimen revealing circular outline and thick pericarp; f: detail of thick pericarp from e; g: detail of innermost pericarp including inner columnar layer (arrows); scale bars: a, c, f, e = 5 mm; b = 10 mm; d, g = 1 mm;

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Plate 22 Lauraceae a–m: Laurocarpum sp. 1, a: SM.B Me 7377, smooth shiny wrinkled cuticle with tiny regularly spaced dimples (? resin cells) on the outer surface; b, c: SM.B Me 13000, fruit in basal and apical views. Note basal scar (in b), and wrinkly surface of cuticle; d: SM.B Me 4993, fruit showing circular basal scar; e, f: M.B Me 2310, fruit in oblique basal and apical views showing large basal scar in e; g: SM.B Me 5112, digital micro CT section, showing a single locule. The thick external cuticle, strongly wrinkled at ends of specimen, is also particularly well seen at right of specimen; h: SM.B Me 2561, fruit showing dark cuticular surface except over the extremely well-defined circular basal scar; i: SM.B Me 5102, basal view of circular scar; j: HMLD Me 1835, fruit with wrinkled surface, oblique side view. This specimen was illustrated as Podocarpus by Engelhardt (1922); k: SM.B Me 4997, oblique side view, with scar comparable to j; l: SM.B Me 5113, oblique side view; m: enlargement of k, showing wrinkled surface cuticle; n: Laurocarpum sp. 2, SM.B Me 4416, fruit identical with Laurocarpum sp. 1 except for being about half the size; note basal scar at arrow; scale bars: a–l = 1 cm; m, n = 5 mm.

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Plate 23 Leguminosae a–f: Mimosites spiegeli Engelhardt, a: type specimen, HLMD-Me-1760, curved pod, with rectangular seed cavities elongate across the pod; b: SM.B Me 7056, similarly curved specimen, seed outline visible in some cavities; c: SM.B Me 19035, stipitate pod with one small calyx lobe on pedicel; d: SM.B Me Me 7057, shorter pod with very clear transverse ridges defining the seed cavities; e: SM.B Me 20054, stipitate pod, seed outlines in basal cavities clearly do not extend to the non-placental margin; f: enlargement of d, showing outline of slightly displaced seed; scale bars: a–e 2.5 cm; f = 1 cm.

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Plate 24 Leguminosae a–d: Leguminocarpon herendeenii sp. n., a: SM.B Me 17519, single seeded pod; b: SM.B Me 21270, single seeded pod with subtending perianth and pedicel, apical extension suggestive of additional undeveloped seed(s); c: holotype, SM.B Me 20422, fruit with two nearly equally developed locules and subtending perianth remains; d: SM.B Me 18271, stipitate fruit with apical locule most developed and apparently opened apically, suggesting dehiscence; e, f: Leguminocarpon sp. 1, e: SM.B Me 7055, nearly straight pod, lacking transverse ridges and with seed development confined mostly to the placental side of the pod. Note rounded-rectangular seed outlines; f: SM.B Me 7035, shorter pod with seeds of similar morphology; g: Leguminocarpon sp. 2., SM.B Me 23628, pod with relatively narrow, transverse seed chambers; h: Mimosites browniana Bowerbank from the London Clay (cementstone lithology), NHMUK V41174, locality of Assington, Suffolk, England for comparison with M. spiegeli (pl. 23d); scale bars: a–d, f–h = 1 cm; e = 5 cm.

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Plate 25 Lythraceae a–f: Decodon-like infructecences, a, b: SM.B Me 2056; a: paniculate infructescence; b: enlargement of a, showing fruits borne on elongate peduncles and remnants of bracts/perianth; c: SM.B Me 17917, fruits showing outlines of seeds; d: SM.B Me 2103, pedicellate fruit with persisting single style; e: SM.B Me 2095, single fruit with cuneiform seeds beginning to detach; f: SM.B Me 7075, two clumps of fruits possibly deriving from a single infructescence. Some of the globose fruits on the right are transversely sectioned, showing the radial arrangement of wedge-shaped seeds; scale bars: a, b = 1 cm; c, d, f = 5 mm; e = 3 mm.

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Plate 26 Lythraceae a–g: Decodon-like, SM.B Me 20157, two adjacent fruits, a: light microscopy showing wedge shaped seed outlines; b–g: digital sections of specimen in a, all of the seeds show dense tissue in outer wall and surrounding embryo cavities and more thin-walled tissue (appearing granular) in remainder of seed coat; b: entire fruit and portion of adjacent fruit showing sections through multiple seeds; c: detail of radially arranged seeds showing various triangular to subquadrangular shapes when sectioned resulting from close packing in the fruit; d, e: planes of section selected to reveal longitudinal sections through seeds showing the slightly curved embryo cavity as a space (black); f: longitudinal section showing detail of dense tissue (no cells discernable, thick walls inferred) and more open thin-walled tissue. Shattered area to right of embryo cavity may represent the germination valve; g: transvere section through seed showing U-shaped embryo cavity (black), shattered tissue between limbs of the U-shape may represent compacted germination valve; h–j: Extant Decodon verticillata Elliott, Alachua Co., Florida for comparison with pl. 25; scale bars: a, h–j = 5 mm, b = 3 mm, d, e = 2 mm, c, f, g = 1 mm.

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Plate 27 Magnoliaceae, Mastixiaceae a–f: Magnolia sp., a: SM.B Me 10611, obovate seed with smooth surface, pointed towards micropylar area; b: SM.B Me 10510, wider seed more rounded towards micropylar area; c: SM.B Me 20156, subtriangular seed; d: SM.B Me 4757, symmetrical obovate seed; e: SM.B Me 7751, transversely elliptic seed; f: enlargement of e, showing seed coat formed of anticlinally oriented columnar cells; g: Magnolia sp. 1., HLMD-Me-15680, multifollicular fruit; h–t: ? Mastixia sp., h: SM.B Me 8359, oblong endocarp with centre of the dorsal furrow occupied by a longitudinal ridge; i: SM.B Me 20574, larger endocarp with obovate shape and surrounded by soft tissue remnant; j: SM.B Me 614, larger endocarp dry specimen showing transverse surface striations (possibly due to drying) and prominent dorsal furrow with a narrow median longitudinal ridge; k: SM.B Me 19992, endocarp with truncate base, posibly damaged and apically flared groove; l: SM.B Me 23368, endocarp showing little compaction and surrounded by soft tissue remnant; m–t: digital SRXTM sections of k; m–o: transverse sections of k, 2/3 from equatorial region to base, 1/3 from equatorial region to base, equatorial region, respectively, no internal compartment (such as a locule or embryo cavity) is evident; p, q: longitudinal sections of k, parallel with surface just below groove, at groove, respectively; some tissue differences are evident within the endocarp wall; r, s: longitdudinal sections of k, perpendicular to surface, through groove and adjacent to groove respectively; t: enlargement of wall in q showing detail of tissue differentiation; more dense areas assumed to contain thick-walled cells; scale bars: a–e, h–l, p–s = 5 mm; g = 1 cm; f, t = 1 mm; m–o = 3 mm.

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Plate 28 Mastixiaceae a–m: Eomastixia cf. rugosa Chandler 1926 emend. Chandler, a: SM.B Me 7154, fruit with longitudinally striate surface; b, c: SM.B Me 20150; b: striate surface with underlying resin pockets revealed in places; c: transverse section of specimen in b, showing that outermost fruit wall contains many large resin pockets; d: transverse section of specimen subsequently used for chemical analysis (van Aarssen et al. 1994); e: SM.B Me 2270, digital SRXTM tranverse section showing resin pockets beneath the surface (darker grey areas); the black irregular lines are cracks and there is no evidence that they have propogated from original locules in the specimen; f: SM.B Me 8895, syncarpous fruit with longitudinally striate surface and apical disk poorly defined; g: SM.B Me 4395, fruit with unusually pointed apex; h: SM.B Me 4127, globose smaller fruit with rounded apex; i: SM.B Me 8267, fruit with base partially fractured, and surface exposing underlying elongate resin bodies; j: SM.B Me 2241, globose fruit in slightly oblique compression; k–m: SM.B Me 20153, four-loculed fruit with U- or V-shaped locules and resin pockets underlying fruit surface, k photographed wet and j dry; m: enlargement of k, showing locule and resin body in outermost fruit wall, SEM; scale bars: a, b, f–l = 1 cm; c–e = 5 mm; m = 1 mm.

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Plate 29 Mastixiaceae a–g: Eomastixia cf. rugosa Chandler 1926 emend. Chandler, a, b: SM.B Me 4053, transversely fractured stone showing five U-shaped locules and large resin pockets underlying fruit surface, a photographed dry, b immersed; c, g: SM.B Me 11280, 3-loculed fruit transverely fractured, g detail of c; d–f: SM.B Me 15448, 2-loculed fruit, transversely fractured; d: transverse fracture; e: transverse section at different level, intercepting a large resin cavity upper right; f: enlargement of d, showing fibrous construction of the endocarp wall; h–l: cf. Beckettia, h: SM.B Me 4118, ovoid endocarp, with a prominent elongate germination valve visible on one face; i: SM.B Me 2663, endocarp, prominent germination valve with a median, longitudinally striate ridge, valve narrows to pointed apex; j: SM.B Me 23139, another specimen with clearly defined germination valve with a median striate longitudinal band; k: SM.B Me 11183, surface smooth, with clear outline of germination valve; l: SM.B Me 17531, more elongate specimen, with clear outline of germination valve; m: NHMUK V23013(3), Beckettia mastixioides from London Clay, for comparison; n, o: SM.B Me 2264, indet. mastixioid; opposite sides of one oblong fruit with prominent longitudinal grooves marking infolds in long, wide germination valves; scale bars: a–d, h–m = 1 cm; e, g, n, o = 5 mm; f = 1 mm.

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Plate 30 Mastixiaceae a–i: Mastixiopsis nyssoides Kirchheimer, a: SM.B Me 5739, fruit in longitudinal fracture showing elliptical outline and thick wall surrounding the elliptical locule with median longitudinal ridge (infold); b: SM.B Me 5082, ellipsoidal fruit with elongate germination valve fallen away revealing thick wall and elongate locule with median ridge; c: SM.B Me 23283, fruit exposed in longitudinal fracture; d: counterpart of the fruit in c; e–i: SM.B Me 7176; e: external view of the fruit; f: reverse side of e from which germination valve has detached; g: germination valve corresponding to the fruit in e, f with an oblique-transverse fracture; h: transverse view of broken piece of the valve from g, showing dorsal infold; i: enlargement from h showing internal anatomy; j, k: Indet. mastixioid fruit, doubtfully conspecific with pl. 29n,o, j: SM.B Me 2225, large broadly ovoid fruit with longitudinal striations and faint longitudinal grooves; k: SM.B Me 2226, large ovoid fruit with faint longitudinal grooves; scale bars: a–h, j, k = 1 cm; i = 3 mm.

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Plate 31 Menispermaceae a–d: Diploclisia rugulosa sp. n., a: SM.B Me 10481, horseshoe-shaped endocarp showing smooth dorsal crest along margin and numerous radiating ribs on the lateral crest; b: holotype, SM.B Me 2069, large endocarp, obliquely obovate in lateral view, showing elongate slit in large elliptic central condylar area; lateral and dorsal crests both ornamented; c: SM.B Me 17772, smaller endocarp showing ornamentation on the lateral and dorsal crests and elongate slit in central area; d: SM.B Me 5155, endocarp where surface ornamentation may be partly obscured by remnants of the outer pericarp; e–h: Stephania hootae sp. n. Endocarps in lateral view showing horseshoe-shaped outline, central foramen, and spines along the lateral and dorsal crests. Shown at same magnification as a–d to emphasize smaller size, e: SM.B Me 5156, showing short spines arranged along the dorsal and lateral crests and protruding from dorsal margin; f: holotype, SM.B Me 2070, endocarp showing one longer limb protruding at base, smooth surface of endocarp surrounding the central foramen and spines on lateral and dorsal crests, protruding form dorsal margin; g: SM.B Me 12253, endocarp with particularly prominent spines along the entire dorsal margin; h: SM.B Me 7583; endocarp with very small spines on the dorsal margin; i: Indeterminate Menispermaceae, SM.B Me 16573; endocarp similar to Diploclisia (a–d above) but less than half the size; j–m: Martinmuellera tuberculata gen. et sp. n., j: SM.B Me 4916, cluster of endocarps of the same species, probably from a coprolite; k: SM.B Me 17404, well-organised cluster of endocarps possibly representing the organisation in an original fruiting branch, tubercles in rows; l: holotype, SM.B Me 8555, isolated endocarp showing distinctive ornament of prominent conical tubercles, irregularly arranged, covering the entire horseshoe-shaped area; m: SM.B Me 4916, detail of one of the endocarps from j; n–r: Endocarps of extant species for comparison, n: Diploclisia affinis, US 599510, Coll E. H. Wilson, China; o: Stephania hernandifolia, US 3345848, Coll A. M. Hug, Bangladesh; p: Stephania delavayi, US 599329, Coll. E. H. Wilson, China; q: Pericampylus glaucus, US 1670049, Coll. C. Wahg, Hainan, China; r: Cocculus carolinus, BARC 19889, Okla, USA; scale bars: a–d, j = 1 cm; e–i, k–r = 5 mm.

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Plate 32 Menispermaceae a: Wardensheppeya sp., SM.B Me 2078. Endocarp with almost symmetrical central condylar area surrounded by lateral crest from which strong radial nodular ridges extend most of the distance across the endocarp, dorsal crest smooth; b–d: Palaeosinomenium ornamentum sp. n. Endocarps with asymmetrical central condylar area, surrounded by a lateral crest from which prominent nodular or club-shaped ridges radiate; between the ends of these ridges and the periphery the endocarp and dorsal crest is finely ribbed, b: holotype, SM.B Me 7343, showing smooth asymmetric central area surrounded by curved lateral crest; short thick club-shaped ridges radiate from the lateral crest reaching about half way to the dorsal margin; c: SM.B Me 2077, endocarp showing elongate foramen, and a row of inconspicuous nodules protruding into the central condylar area from the lateral crest; d: SM.B Me 7212, similar to c but lacking nodules on the condylar side of the lateral crest and having very clear fine ribbing on the dorsal endocarp; e: Palaeosinomenium venablesii Chandler, SM.B Me 14061, very small endocarp with asymmetric central condylar area and prominent radial ridges extending from the lateral crest but mostly not to the endocarp periphery; f–i: Palaeosinomenium sp.; endocarps with strongly asymmetrical base, asymmetrical central condylar area surrounded by a lateral crest from which narrow, fine, simple short ridges radiate, usually continuing across the dorsal crest and sometimes protruding at the periphery, f: SM.B Me 4011, endocarp with nodules protruding from the lateral crest into the central condylar area; radial ridges extend from lateral crest across endocarp onto dorsal crest and protrude from the periphery; g: SM.B Me 4193, endocarp in which radial ridges do not protrude at periphery; h: SM.B Me 2073, small example with very fine ridges extending to, but not beyond, periphery; i: SM.B Me 17855, endocarp with strongly asymmetrical base and prominent elongate foramen in the central condylar area; j: cf. Pericampylus, SM.B Me 23272. Endocarp with oblique base and very small size, dorsal crest with prominent long dorsal spines, and a row of shorter spines extending from the margin of the lateral crest; k–r: Cocculus lottii sp. n. Endocarps lacking dorsal crest but with a groove marking the plane of bisymmetry in an otherwise broad, rounded dorsal surface; with small central depression on each lateral face surrounded by an annular bulged area with radiating rugulae, k: SM.B Me 4152, endocarp with prominent sediment filled central depression from which coarse rugulae radiate, prominent groove in plane of bisymmetry; l: SM.B Me 7187, endocarp with central depression, coarse rugulae and deep groove in plane of bisymmetry cleary displayed by oblique compression tilted towards viewer; m: SM.B Me 4242, endocarp with fine rib-like rugulae; n: SM.B Me 2228, endocarp with oblique compression, so that broadly rounded dorsal surface is visible at right of image. Some rugulae elevated into spine-like ornament protruding from endocarp surface; o: holotype, SM.B Me 8559, endocarp with central depression from which coarse rugulae radiate, compressed such as to tilt the ventral surface towards the viewer; groove in plane of bisymmetry visible at lower left of specimen; p: SM.B Me 7214, endocarp compressed to display rounded doral surface with prominent groove; q: SM.B Me 13030, endocarp cluster; r: SM.B Me 20562, pedicellate smaller specimen, possibly belonging to a second species; s–v: Endocarps of extant species for comparison, s: Menispermum dauricum, BARC 61791; t: M. canadensis, UWM15599, image courtesy Sara Hoot; u: M. canadensis, Harrisonburg, Virginia, USA; BARC 94DR; v: Sinomenium acutum, Szechwan, China, BARC 114802; scale bars: a–j, r–v = 5 mm; k–q = 1 cm.

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Plate 33 Menispermaceae a–o: Karinschmidtia rotulae gen. et sp. n., a: SM.B Me 2001, circular fruit, with pericarp outer cuticle and concentric horseshoe shaped ridges of the endocarp evident in the centre; b: SM.B Me 7037, fruit with pericarp cuticle partially freed from sediment, casting shadow; c, m: holotype, SM.B Me 21233, circular fruit showing radiating strands extending distally from the ridged endocarp; d, e: SM.B Me 20452, tough radiating strands with terminal branching in the circumference, smooth central portion (concave) of endocarp; f: SM.B Me 17153, showing cuticle sheath extending beyond the margins of the endocarp wing; g: SM.B Me 8553, detail of fibrous strands of the endocarp wing; h: SM.B Me 18873, isolated endocarp missing its wing; i: SM.B Me 20543, fruit with clearly defined endocarp crests (dorsal and paired lateral); j: SM.B Me 13487, fruit photographed in dry condition by reflected light; k: specimen in j, photographed with low magnification epifluorescence to show cuticle; l: specimen in j, under higher magnification epifluorescence, cuticle showing a polygonal pattern of epidermal cells; m: SM.B Me 21233, enlargement from c showing distinct fibrous strands perpendicular to fruit margin; n: SM.B Me 24310, basal half of a fruit showing radially striate endocarp wing and pericarp cuticle extending beyond the fibrous strands also showing the longer stylar limb and shorter hilar limb of the endocarp, the latter opposite the likely attachment position; o: SM.B Me 24310, counterpart of n, apical half showing cuticle extending beyond the margin of the fibrous strands; p–r: Legnephora minutiflora (K. Sch.) Diels, extant endocarp for comparison, northeastern New Guinea, Arnold Arb. Herbarium: Clemens 8682, p: lateral view showing partially encircling winglike dorsal crest and single (rather than double) lateral crest; q: dorsal view showing wide central area with protruding lateral cretsts, and narrow, winglike median dorsal crest; r: detail of fibrous wing-like dorsal crest by combined reflected and transmitted light; scale bars: a–d, i–k, n, o–q = 1 cm; e–h, m, r = 5 mm; l = 150 µm.

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Plate 34 Menispermaceae, Myristicaceae a–c: Tinomiscoidea jacquesii sp. n., a: holotype, SM.B Me 2066, ventral side of endocarp showing pointed apex, median longitudinal groove and transversely to slightly obliquely oriented striae diverging from the median line; b: SM.B Me 2068, endocarp in dorsal view, finely verrucate, with wide median and marginal ridges reflecting collapse of the dorsal wall conforming to the ventral topography; c: SM.B Me 2067, dorsal side, showing pointed apex, rounded base, and finely verrucate surface; d–f: Parabaena cf. europaea Czeczott & Skirgiełło, d: SM.B Me 13741, specimen viewed dorsally with median keel transverse striate patterning, and peripheral spines; e, f: SM.B Me 4023, another specimen showing protrusion of spines around periphery; g, h: Tinosporeae sp. 1., g: SM.B Me 2216, ventral surface of endocarp bisymmetrical about a median dorsal ridge, smooth; h: SM.B Me 2218, dorsal surface of endocarp bisymmetrical about a median dorsal ridge, coarsely veruccate to rugulate; i, j: Tinosporeae sp. 2, SM.B Me 2213. Dorsal and ventral view of smooth endocarp; k: Unnamed Menispermaceae, SM.B Me 4091, strongly curved endocarp with three spiny crests and ridges radiating from the condylar depression (seen at right); l–n: Tinosporeae sp. 3, l: SM.B Me 2187, dorsal view of endocarp ruptured apically, showing sharp keel and verrucate surface; m: SM.B Me 8830, lateral view with convex dorsal margin to left, showing verrucate surface and flattened ventral surface; n: ventral view of m showing central depression and apical keel; o, p: Myristicacarpum sp., o: SM.B Me 5605, seed circular in profile; p: same specimen as in o viewed on transverse fracture plane. Ruminate endosperm with irregular longitudinal plate-like intrusions of the seed coat; scale bars: a–c, o = 1 cm; d–n, p = 5 mm.

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Plate 35 ? Nymphaeales a–i: ? Nymphaeales, unnamed seeds, a: SM.B Me 2735, large cluster of one kind of seed; b: SM.B Me 7011, small cluster with some specimens apparently loose suggesting that they were originally held together by mucilaginous material; c: SM.B Me 7014, seeds from a loose cluster, showing longitudinal raphe ridges; d: SM.B Me 7666, isolated seed with apical operculum and raphe along left lateral margin; e, g–i: SM.B Me 4045; e: SEM of seed showing sub-basal chalaza (seen as a round protrusion) and an operculum (terminal circular apical cap); f: SM.B Me 7012b, SEM of operculum showing polygonal cells, circular central hilum and micropylar slit; g: enlargement from e, operculum (arrow marks its margin) with protruding hilum; h: detail from e showing surface of the sclerotic seed coat consisting of interlocking digitate cell outlines with micropapillate to micropitted surface; i: detail from h showing seed surface minutely papillate and pitted; scale bars: a, b = 1 cm; c = 2 mm; d = 3 mm; e = 1 mm; f = 150 µm, g = 100 µm; h = 50 µm; i = 10 µm.

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Plate 36 Nyssaceae, Pentaphylacaceae, Rhamnaceae a–d: Nyssa disseminata (R. Ludw.) Kirchheimer, a: SM.B Me 7566, endocarp showing elliptical outline and curved outline of apical germination valve; b: SM.B Me 7207, endocarp with elliptic outline to germination valve, arched slit around valve confined to upper half of the fruit, and shallow longitudinal ribs; c: SM.B Me 7194, endocarp with somewhat more pointed base and apex. Note outline of germination valve in the apical 1/3; d: SM.B Me 8256, more elongate specimen showing surface pattern of low meridional ribs; e–g: Cleyera sp., e: SM.B Me 12164, campylotropous seed with shallow truncated micropyle and chalazal ends; f: SM.B Me 20158, showing shallow median depression around which the C-shaped embryo cavity curves; g: detail of f, showing reticulum of polygonal cell outlines with upraised outer periclinal walls; h–n: Berchemia mellerae sp. n., h: SM.B Me 8793, endocarp with a smooth surface with a distinct narrow median groove and truncate base; i: SM.B Me 4773, more narrowly elongate example; j: SM.B Me 19004, endocarp retaining a portion of apparent outer softer tissue at upper right; k: SM.B Me 21298, shorter example; l–n: holotype, SM.B Me 4184; l, m: endocarp released from oil shale to show both surfaces with median ridge continuous around the endocarp; n: basal view, showing lensoidal outline with median ridge; o: Extant Berchemia lineata DC. A. E. H. Wilson 11080, Taiwan for comparison; scale bars: a–d, h–k = 1 cm; e, f = 4 mm; g = 2 mm; l–n = 5 mm, o =3 mm.

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Plate 37 Rutaceae a–e: Rutaspermum messelense Collinson & Gregor, a: SM.B Me 17137, seed showing surface ridges and flattened apical portion of ventral face; b–e: SM.B Me 7952, seed released from oil shale shown in various views with surface of longitudinal ridges with anastomoses, flattened apical portion of ventral face with elongate triangular hilar scar, seed surface around hilar scar slightly inflated and more or less smooth; f: Rutaspermum undetermined A., SM.B Me 4037, of Collinson & Gregor (1988) showing longitudinal ridges lacking anastomoses; g–i: Rutaspermum undetermined B. of Collinson & Gregor (1988), g, h: SM.B Me 7372, lateral and ventral view; showing reticulate surface lacking prominent ridges, hilar scar extending almost full length of ventral face and raphal excrescence, probably belonging to this taxon; i: SM.B Me 2587, specimen attributed to this taxon by Collinson & Gregor (1988), with surface reticulum; j–l: cf. Toddalia sp., j: SM.B Me 4142, seed with an elongate-elliptical raised hilar scar almost terminal on the ventral face and possible median dehiscence slit; k: SM.B Me 4677, smooth surface, finely pitted with a pattern of polygonal cells; l: SM.B Me 4304, with raised hilar scar; m: Rutaspermum chandleri Collinson & Gregor, SM.B Me 4132. Seed with smooth surface and flat hilar face; scale bars: a–m = 5 mm.

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Plate 38 Sabiaceae, Sapotaceae, Simaroubaceae a–f: Meliosma sp., a, b: SM.B Me 10620, endocarp with keel in plane of symmetry, ventral depression with prominent funicular cavity or plug; c: SM.B Me 8818, endocarp with prominent ventral depression; d: dorsal surface from c showing median keel; e, f: SM.B Me 21304, strongly compressed endocarp, in oblique-ventral (e) and oblique dorsal (f) views; g–i: cf. Meliosma sp. SM.B Me 4111, small specimen possibly representing a distinct species; j, k: Sapotispermum sp., SM.B Me 8333, j: elliptical seed showing pronounced oblique truncation; k: detailed enlargement showing thin cuticle layer remaining over patches of the seed coat; l, m: Ailanthus confucii Unger, l: SM.B Me 4006, small biwinged fruit with central seed; m: SM.B Me 21808, larger specimen more typical of the species; scale bars: a–f, j, l, m = 1 cm; g–i = 5 mm; k = 1 mm.

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Plate 39 Tapisciaceae, Toricelliaceae a–l: Tapiscia pusilla (Reid & Chandler) Mai, a: SM.B Me 13060, fruit in ventral view showing ovate outline, veruccate-rugulate outer layer of fruit surface, and central concavity marking underlying chalaza; b: SM.B Me 12243, smaller specimen, ventral view; c: SM.B Me 12234, relatively large specimen, ventral view; d: SM.B Me 2246, additional example; note apical bulge at micropyle; e: SM.B Me 24289, large cluster of fruits, assumed to be from a coprolite; f: SM.B Me 17741, fruit abutting an elongate cuticular structure with widened tip, possibly swollen pedicel; g, h: SM.B Me 2189, ventral and dorsal surfaces of a compressed fruit, removed from shale; note that dorsal surface bears no particular distinguishing features and could be confused with other taxa with similar ornament; i: SM.B Me 21386, specimen in oblique, lateral view showing typical verrucate rugulate surface; main diagnostic characters hidden within sediment; j–l: same specimen as in i, digital sections by SRXTM; j: digital oblique longitudinal section, parallel to surface, showing smooth surface of seed, elongate sediment–filled chalazal depression, and apical micropylar bulge; k, l: partial digital sections at slightly different depths to j showing smooth seed surface and, in l especially, distinction between denser (darker) seed tissue and surrounding less dense (lighter) fruit tissue; m–p: Toricellia bonesii (Manchester) Manchester, m: SM.B Me 12260, specimen in dorsiventral compression, ventral view, showing key diagnostic characters. Two main ridges either side of the seed chamber and the subtriangular germination valve are clearly visible. In this specimen a median ridge extends the full length of the seed chamber. [in contrast, in SM.B Me 5745, the median ridge is only present on the germination valve]; n: SM.B Me 4622, specimen compressed laterally, poorly preserved and with some damage during preparation; o: SM.B Me 2197, specimen compressed laterally, D-shaped profile with ventral ridges encircling two-thirds of the endocarp (right of image) and one lateral-dorsal hole in endocarp wall (left of image); p: SM.B Me 2198, specimen in lateral compression. One lateral-dorsal endocarp hole is clearly visible; scale bars: a–d, f–p = 5 mm; e = 1 cm.

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Plate 40 Theaceae a–i: Camelliacarpoidea messelensis sp. n., holotype, SM.B Me 8953, a: subglobose fruit, finely rugulate, apparently leathery; with circular basal disk subtending persistent calyx; b: enlargement of a, showing persistent hypogynous calyx with four imbricate lobes clearly seen and a fifth inferred from symmetry; c: enlargement of b, showing brittle cracked, crushed elongate seed with a shiny seed coat and indication of polygonal surface pattern; d–i: SEM of a very small fragment removed from c; d: seed coat fragment with remnants of possible outer non-sclerified thin tissue layer; e: detail of c, showing surface of seed coat with polygonal cell outlines each polygon with an indistinct central pit; f: broken edge of the seed, showing surface cells, underlying columnar cells and thin inner folded cuticular layer; g: very thick-walled columnar sclereids of seed coat with small lumina, decreasing in size outwards; h: enlargement of g, showing lumina and angular facets conforming to surface cell shapes; i: detail of pitting inside lumen of sclereid; scale bars: a, b =1 cm; c = 3 mm; d = 200 µm; e, g = 50 µm; f, h = 20 µm; i = 5 µm.

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Plate 41 Ulmaceae a–f: Cedrelospermum leptospermum (Ettingshausen 1853) Manchester 1987 emend. Wilde & Manchester a: SM.B Me 15393, single axillary fruit attached to a zig zag twig with one leaf; b: multiple fruits attached to a leafy shoot, specimen missing; c: SM.B Me 15974, multiple fruits attached to a leafless twig; d: detail of fruits from b; e: SM.B Me 7274, isolated fruit; f: detail from c, paired fruits with veins converging distally and evident distal stigmatic notch; scale bars: a–c, f = 1 cm; d, e = 5 mm.

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Plate 42 Vitaceae a–h: Vitis messelensis sp. n., a–h: Four seeds released from oil shales to show both surfaces; ventral surfaces show narrow infolds parallel at base and then diverging apically whilst dorsal surfaces show pyriform (f) to elliptical chalaza; all surfaces are smooth and all specimens show prominent basal truncate beak. Apical notch shallow (e, f) to missing; a, b: holotype, SM.B Me 21619, ventral, dorsal; c, d: SM.B Me 4025I, ventral, dorsal showing slight fluting radiating from chalaza; e, f: SM.B Me 4025II, ventral showing raphe with slit-like groove, dorsal; g, h: SM.B Me 7282, ventral, dorsal; i–k: Parthenocissus britannica (Heer) Chandler, i, j: SM.B Me 12271, seed with ventral infolds (i) linear, slit-like, straight, relatively long; k: SM.B Me 7337, seed in oblique lateral view; elliptical chalaza near right; l: Vitis messelensis sp. n., SM.B Me 7123, two seeds within same fruit; m–r: Parthenocissus britannica (Heer) Chandler, m: SM.B Me 7974, seed showing relatively long divergent ventral infolds; n: seed in m removed from shale, and turned to show dorsal surface with elliptical chalaza; note absence of apical groove; o: seed in m, fractured edge showing thickness of seed coat; p: SM.B Me 5744, fruit showing two seeds; q, r: SM.B Me 15840, cluster of seeds with remnant of pericarp; scale bars: a–n, p, r = 5 mm; o = 1mm; q = 1 cm.

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Plate 43 Vitaceae a–i: Ampelopsis sp., a: SM.B Me 2339, seed in dorsal view with broad ridges radiating from a centrally positioned chalaza; b: SM.B Me 17395, seed in ventral view showing broadly triangular outline, widely divergent ventral infolds and prominent raphe ridge; c–f: Two seeds released from oil-shale to reveal both surfaces, both surfaces show radiating ridges, those on the dorsal surface being particularly pronounced and sharp-crested; c, d: SM.B Me 20381; e, f: SM.B Me 2352; g: SM.B Me 5166, impression of chalaza and radiating ridges in shale with only a portion of the the original seed tissue remaining on right side of impression; h, i: enlargements from g, showing thickness of seed coat fractured showing columnar arrangement of cells; j–n: Palaeovitis sp., j, k: SM.B Me 7638, seed removed from oil shale to show both surfaces; almost circular in outline, dorsal surface with spatulate chalaza, ventral surface with median ridge but details of infolds obscured by damage; l: SM.B Me 10676, seed showing radiating dorsal ridges; m: SM.B Me 4936, another specimen with large spatulate chalaza; n: detail from k, through portion of fractured seed coat showing isodiametric, rather than columnar cells; o, p: Cayratia jungii (Gregor) Chen & Manchester, SM.B Me 7613, seed removed from oil shale to show both surfaces, seed elliptical with subparallel sides, o: dorsal surface with narrowly elongate chalaza and radiating ridges; p: ventral surface with shallow infolds; q, r: Unnamed Vitaceae; radially striated seed, SM.B Me 7983, q: ventral view showing long, apically divergent ventral infolds and striated surface; r: dorsal view with crushed surface with inconspicuous chalaza and irregular striations; scale bars: a–g, j–m, o–r = 5 mm; h = 2 mm; i, n = 0.5 mm.

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Plate 44 Vitaceae a–m: Crassivitisemen wildei (Chen & Manchester) comb. n., a: SM.B ME 2298, fruit in oblique lateral view, showing prominent longitudinal vascular strand; b–d: SM.B Me 5727, fruit in ventral, dorsal, and polar view; e: SM.B Me 8410, fruit in shale, fractured transversely; f–h: internal detail from a, revealed by digital SRXTM sections; f: transverse digital section showing seed with prominently protruding raphe ridge separating two ventral infolds, the entire seed enveloped by pericarp which has a dense inner layer and thick outer layer of larger cells; g: partial longitudinal digital section of specimen showing very thick seed coat, with ridges on its outer surface, and the pair of longitudinal ventral infolds; h: detail of seed coat and fruit wall anatomy seen in digital longitudinal section (at right angles to k), from left to right, thick outer pericarp with thin-walled, large-celled parenchymatous tissue, thinner inner pericarp with dense tissue of thicker-walled cells, thick seed coat composed of compact anticlinally arranged rows of small nearly isodiametric cells. At the furthest right are remnants of decayed tissue from within the seed; i: fractured transverse surface from e, showing single thick-walled seed within fruit; j–m: SM.B Me 5729; j: isolated seed in dorsal view showing rugulate surface; k: physical transverse section showing prominent ventral raphe ridge; l: enlargement from k, showing extremely thick wall of the seed; m: detail from k. Note nearly isodiametric cells in compact anticlinal rows composing the seed coat, and surrounding darker tissue of the pericarp; scale bars: a–e, j = 1 cm; f, g, k, i = 5 mm, h = 1 mm; i = 0.5 mm; l = 2 mm; m = 1 mm.

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Plate 45 Vitaceae, Carpolithus a–d: Crassivitisemen wildei (Chen & Manchester) comb. n., a: SM.B Me 8786, dorsal surface with ridges radiating out from the edge of the elongate oval chalaza; b: SM.B Me 8205, ventral surface with slightly divergent linear infolds; c, d: holotype, SM.B Me 7271, in ventral and dorsal view; e–l: Carpolithus callosaeoides (Engelh.) comb. n., e, f: HLMD-Me-5333. [Carya costata Unger sensu Engelhardt, 1922, pl. 32, fig. 12]; e: widely ovate fruit, surface with longitudinal striations; f: rounded base with a prominent bulging circular scar; i, j: HLMD-Me-2142. [Ficus callosaeoides type, Engelhardt, 1922, pl. 12, fig. 6a,b]; i: fine longitudinal striations radiating from prominent basal scar; j: reverse side; g, h, k, l: SM.B Me 2322; g: ovate fruit, side view, with prominent basal scar on oblique lower surface; h: same, reverse side view, with apical depression visible; k: cross section showing thick wall of fruit and single-seeded locule; l: enlargement of k, showing the elongated resin or latex-filled cells in cross section; scale bars: a–d = 1 cm; e–j (same bar applies) = 2 cm; k = 5 mm; l = 3 mm.

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Plate 46 Saportaspermum and Spirellea a–h: Saportaspermum kovacsiae Kvaček & Wilde, winged seeds, a: SM.B Me 5576, ovate seed body with tapering lateral wing; b: SM.B Me 21809, elliptical seed body and relatively wide wing; c: SM.B Me 8586, wing with straight upper margin and convex lower margin; d: SM.B Me 21810, seed body showing a faint longitudinal striation; e: SM.B Me 5587 (holotype), showing oblique angle between long axis of wing and long axis of seed body; f: SM.B Me 20349, seed body showing a relatively large circular basal scar; g: SM.B Me 17447, slender specimen with evident basal scar; h: detail of g showing a basal circular scar, and wing lacking venation; i–m: Spirellea sp., SM.B Me 2196, i: specimen as exposed in shale; j–m: same, removed from shale; j: specimen showing longitudinal ridges meeting at both ends, and transverse striae; k: specimen rotated 180 degrees, showing similar ornamentation; l: end view with a smooth elliptical scar; m: opposite end to l showing, less ornamented conical tip; scale bars: a–g = 1 cm; h–m = 5 mm.

186


Plate 47 a–f: Carpolithus sp. 1, a: SM.B Me 7334, fruit showing basal scar with raised rim and meridional grooves; b: SM.B Me 7095, laterally compressed fruit showing pyriform shape and parts of five longitudinal valves; c: SM.B Me 19836, laterally compressed fruit showing prominent basal scar and rounded apex; d: SM.B Me 18096, laterally compressed fruit showing smooth, somewhat shiny surface, and separation of valve at apex; e, f: SM.B Me 15049, partially disaggregated fruit in lateral view; f: detail of valve surface, smooth appearance but densely verrucate at high magnification; g–l: Carpolithus sp. 2, g: SM.B Me 2211, typical specimen showing elliptical body and long narrow style; h: SM.B Me 17776, specimen with very long style preserved; i: SM.B Me 7605, specimen with well preserved seed body, with a small apical scar; j, k: SM.B Me 17776, successive enlargements, showing a halo of tissue that surrounds the elliptical body and is continuous with the style, small circular scar at apex of seed body; l: SM.B Me 2211, enlargement, showing longitudinally striate elliptical body; scale bars: a–e, g, h = 1 cm; f, j = 5 mm; i = 6 mm; k, l = 3 mm.

188


Plate 48 a–i: Carpolithus sp. 3, a: SM.B Me 7070, obovate fruit showing fibrous wall at base; b: SM.B Me 7068, surface coarsely transversely corrugated; c: SM.B Me 7065, external surface of fruit partially broken at base showing moderately thick wall; d, e: SM.B Me 2325, fruit removed from shale, viewed from both sides; f: SM.B Me 8807, three closely associated fruits; g: SM.B Me 13738, single fruit showing acute apex and somewhat broader base; h: SM.B Me 8807, enlargement from f, showing simple black, unbranched hairs; i: SM.B Me 7065, enlargement from c, showing simple black, unbranched hairs; j–n: Carpolithus sp. 4, j: SM.B Me 21562, ovoid seed, undulating surface suggesting ductile (not brittle) deformation during compression; k: SM.B Me 5057, globose specimen; l: detail from k, showing striate appearence resulting from rectangular surface cells in longitudinal rows; m, n: detail of j, showing rectangular surface cells aligned in longitudinal rows; scale bars a–g = 1 cm; h, i = 1 mm; j, k = 5 mm; l–n = 0.5 mm.

190


Plate 49 a–c: Carpolithus sp. 5, SM.B Me 5555, a: inside surface of fruit with smooth margins, undulate locular area and central longitudinal fibre bundle; b: same, enlarged, showing compressed tissue of the locular area, possibly including winged seeds; c: detail from b, showing three fragmentary overlapping seeds; d–g: Carpolithus sp. 6, d: SM.B Me 4114, globose pedicellate fruit with truncate base and elliptical black shiny seed; e: SM. B Me 4125, pedicellate fruit apparently wider than long, outer cuticle preserved, softer inner tissues lost, containing several dark shiny seeds; f: SM. B Me 13028, fruit slightly damaged, containing two or more seeds (arrows); g: detail from f showing dark shiny elliptical seed; h–k: Carpolithus sp. 7, h: SM.B 8539, thick pedicel, oblate capsular fruit with meridional grooves, and one of the styles persisting, arching to the left of the apex; i: SM.B 8057, pedicellate capsule with two recurved styles visible at apex; j: SM.B 14666, apical portion of fruit, obliquely oriented, with two very obvious persistent styles; k: SM.B 8037, fruit showing finely verrucate texture and meridional grooves delimiting valves; scale bars: a, b, d–f = 1 cm; c = 2 mm; g = 3 mm; h–k = 5 mm.

192


Plate 50 a–c: Carpolithus sp. 8, SM.B Me 15860, a: grouping of angular pyrenes/nutlets/seeds; b: detail of a showing irregularly dimpled face and smooth margins; c: detail of b showing surface pattern of fine cells forming longitudinal striae in the center and transverse striae over corner ridges; d, e: Carpolithus sp. 9, SM.B Me 16405, d: specimen as exposed in shale with a median rib separating two concavities and surrounding softer tissue; e: removed from shale, and turned over to show opposite surface from d; f: Carpolithus sp. 10, SM.B Me 13973, five-seeded berry, with cuticle indicating inner collapsed soft tissue; g: Carpolithus sp. 11, SM.B Me 5586, transversely compressed specimen showing central depression surrounded by ca 10 wedge shaped units separated by grooves; scale bars: a, f, g = 3 mm; b = 1 mm; c = 0.5 mm; d, e = 5 mm.

194


Plate 51 a–h: Carpolithus sp. 12, ? Ericales, a: SM.B Me 10447, pedicellate fruit attached to portion of a branched infructescence; b: SM.B Me 7096, pedicellate, obovate fruit in lateral surface view with longitudinal ridges, hypogynous calyx and stylar protrusion; c: SM.B Me 7235, dehiscent fruit showing basal perianth scar, axial vascular strand and two attached elliptical seeds; d: SM.B Me 7591, isolated seed with finely papillate surface; e: SM.B Me 7235, detail from c showing finely papillate, in situ seeds; f: SM.B Me 10447, detail from a showing outline of seed within the fruit; a fragment of the fruit removed, revealing papillate surface of the seed within (arrow); g: SM.B Me 8702, isolated elliptical papillate seed; h: SM.B Me 8702, detail from g showing papillate surface; i, j: Carpolithus sp. 13, SM.B Me 7064, i: globose fruit with stout pedicel; j: detail from i showing prominent disk at base of fruit, smooth locule surface and arched central strands indicating possible site of placentation; scale bars: a–c, j = 1 cm; d, e, g, h = 5 mm; f = 3 mm; i = 2.5 cm.

196


Plate 52 a–f: Carpolithus sp. 14, SM.B Me 4086, Sambucus-like seed cluster, a: cluster of seeds within fruit; b: detail from a showing two overlapping seeds with corrugated surfaces; c: single seed from a, showing corrugated surface; d: same seed as c, showing low topography of surface ornamentation, SEM; e, f: detail from d of thick-walled isodiametric cells at the seed surface, SEM; e: surface cells retaining their outer periclinal walls; f: in an area where surface cells are abraded and cell lumina are visible; scale bars: a = 5 mm; b, d = 200 µm; c = 100 µm; e, f = 10 µm.

198


Plate 53 a–d: Carpolithus sp. 15, HLMD-Me-13399, a: fruit with meridional grooves and surface ornamented with evenly distributed, short, blunt spines; b: detail from a showing longitudinal groove separating two valves; c, d: details from a showing short, broad-based blunt spines; e–g: Carpolithus sp. 16, SM.B Me 11013, e: branched infructescence with at least three orders of branching; f: detail of elliptical fruits with widely spaced longitudinal striae; g: fruit with subtending perianth scar (arrow); scale bars: a = 2.5 cm; b, e = 1 cm; c, f, g = 5 mm; d = 4 mm.

200


Plate 54 a–d: Carpolithus sp. 17, SM.B Me 2733, a: branched infructescence with numerous ovoid to globose fruits; b, c: details from a showing swollen pedicel, perianth scar at the base of the fruit and apical circular bulge; d: enlargement of a showing branched infructescence with bracts at nodes (arrows) and non-attached leaves; scale bars: a, d = 3 cm; b, c = 1 cm.

202


Plate 55 a, b: Carpolithus sp. 18, HLMD-Me-16934, a: fruit with sinusoid notch (? damage) at one end, split longitudinally showing thick wall, smooth locule and multiple medially positioned seeds; b: detail of a showing large shiny seed; c, d: Carpolithus sp. 19, SM.B Me 17403, c: pedicellate, widely ovate fruit with disk like structure at base of fruit; d: detail of c showing a single obovate seed within margin of fruit although not conclusively belonging to the fruit; note free membranous edge (arrow) possibly representing a wing; e, f: Carpolithus sp. 20, SM.B Me 8404, e: pedicellate, fin-winged fruit with at least four wings; f: enlargement showing overlapping margins of two wings, with strong fimbrial veins and relatively widely spaced transverse veins (arrows); g, h: Carpolithus sp. 21, g: SM.B Me 10508, aggregation of numerous tiny elliptical winged seeds; h: SM.B Me 8708, elliptical seed with transverse and longitudinal ridges surrounded by a smooth membranous wing; scale bars: a–f = 1 cm; g = 3 mm; h = 1 mm.

204


Plate 56 a–k: Carpolithus sp. 22, a: SM.B Me 484, ellipsoidal fruit with numerous stout spines arranged in longitudinal rows; b: SM.B Me 2059, fruit with numerous spines, a few of which are distally bifurcate; c: HLMD-Me-13925, fruit with ridges bearing stout spines; d: HLMD-Me-16935, broken end of fruit exposes an underlying smooth surface with a median groove; e–k: SM.B Me 2059, successive digital transverse sections, SRXTM series showing unilocular construction, and thick wall with spines protruding in all directions, four main rows of spines, (left and right and median on either side); l, m: Carpolithus sp. 23, l: SM.B Me 4174, fruit composed of three radially arranged bipartite units; m: SM.B Me 10588, fruit with prominent recessed area at one end from which it is splitting; n, o: Carpolithus sp. 24, SM.B Me 17400, n: elliptical seed with a slightly sinuous median longitudinal groove and transverse ridges; o: detail from n showing shiny cuticle-testa layer adhering in patches; p: Carpolithus sp. 25, SM.B Me 7352. Fruit with meridional veins separating each carpel; q–s: Carpolithus sp. 26, tufted disseminule, q: SM.B Me 20154, elliptical seed/fruit with numerous hairs arising from base; r: SM.B Me 23128, specimen showing smooth elliptical body and attached hairs; s: SM.B Me 23128, detail from s, showing overlapping but unbranched hairs; scale bars: a–d = 1 cm; e–n, p = 5 mm; o, s = 2 mm; q, r = 3 mm.

206


Plate 57 a–d: Carpolithus sp. 27, SM.B Me 7040, a: raceme bearing numerous, nearly sessile fruits; b: detail of a, showing fruits with persistent tepal-like lobes arising from rim of hypanthium; c: detail of a, showing peglike, very short broad pedicels and fruits with widely spaced longitudinal ribs, one with tepal-like lobes open, the other closed; d: detail of a, showing peg-like pedicel, longitudinal ribs on the hypanthium and fine venation and wrinkles on the perianth lobes; e–g: Carpolithus sp. 28, SM.B Me 21351, e: elliptical disseminule, with a prominent circular or elliptical basal scar, photographed immersed in water; f: photographed partially dry; g: SRXTM digital transverse section showing single flattened central cavity and thick wall with anticlinal cracks; scale bars: a = 3 cm; b–d, g = 5 mm; e, f = 1 cm.

208


Plate 58 a–g: Carpolithus sp. 29, a: SM.B Me 7190, infructescence bearing several elliptical to obovate fruits; b: detail of a, showing prominent disk at the top of the pedicel immediately below each fruit; c: SM.B Me 7540, infructescence with attached capsules; d: SM.B Me 7218, infructescence with obovate apically truncate fruits; e: SM.B Me 15411, infructescence with three branches arising from main axis; f: detail of e, showing fruits with prominent basal disks, and a dehiscent fruit showing the separation of three valves (arrows); g: SM.B Me 17210, two fruits of a single infructescence oriented to show three valves (left) and two of the valves (right). Note doubly thickened disk at base of fruit (arrow); h, i: Carpolithus sp. 30, SM.B Me 20155, h: elliptical fruit with a distinct rounded stylar protuberance; i: detail of area with part of the pericarp missing, exposing numerous subglobose seeds with a circular depression at one end; scale bars: a–i = 1 cm.

210


Plate 59 a–k: Carpolithus sp. 31, a: SM.B Me 4030, seed with deep cleft; b: SM.B Me 5148, non-ridged side of seed; c–e: SM.B Me 4029; c: seed in shale, showing narrow ridge running from the base of the cleft to the broader end of the seed; d: same, removed from shale, showing ridged face of seed; e: same, reverse side, showing non-ridged face of seed; f: SM.B Me 4016, complete elliptical specimen with typical apical cleft; g: SM.B Me 17234, larger specimen with narrow median ridge and deep cleft; h: SM.B Me 17294, seed showing narrow ridge, and bulging and thickened asymmetrical cleft, photographed dry; i: detail from h, showing cellular pattern at surface; j, k: SM.B Me 2166, SEM of seed fragment showing columnar construction; l: Carpolithus sp. 32, SM.B Me 7079, m–q: Carpolithus sp. 33, SM.B Me 12158, m: pedicellate immature fruiting head with apical extensions to immature fruits in the upper portion; n–q: micro CT digital sections; n: longitudinal section through pedicel and showing at least one fruit extension protruding into oil shale; o–q: top, middle, and bottom transverse sections, respectively; showing fruits and a few of their apical extensions protruding into the oil shale, especially in the lower part; scale bars: a–h, n–q = 5 mm; i = 3 mm; j = 400 µm; k = 200 µm; l, m = 1 cm.

212


Plate 60 a–l: Carpolithus sp. 34, a: SM.B Me 16388, seed with surface reticulum; b: SM.B Me 4782, large specimen; c: SM.B Me 2089, specimen with obvious reticulum; d: SM.B Me 7100, more pointed specimen; e: SM.B Me 7092, three seeds enclosed in a weak circular structure; f: detail of e showing, one well defined seed; g–l: SM.B Me 4782, SEM of broken seed from b; g: broken longitudinally showing single internal cavity and beveled edges; h: vertical section of seed coat, wall not columnar; i–l: surface of seed; i: showing pattern of fine polygonal pits; j: outer layer of polygonal cells (lower part of image), partially removed and revealing underlying layer (upper part of image; see k); k: subsurface layer of thick-walled cells with sunken quadrangular lumina, possible position of crystals; l: detail of outer part of h, showing outer thick-walled cells, inner anatomy unclear; m–o: Carpolithus sp. 35, SM.B Me 23088, m: cluster of at least four sessile, seeds or endocarps; n: detail of m, showing longitudinal venation; o: detail of n, showing thin loose wrinkled tissue (bottom left) that may envelope all the seeds/endocarps; scale bars: a–f = 5 mm, g= 2 mm; h = 100 µm, i = 300 µm, j =50 µm, k, l = 20 µm; m, n = 1 cm; o = 3 mm;

214


Plate 61 a–1: Carpolithus sp. 36, a: SM.B Me 7619, isolated head showing fibrous central core and outer compartments at left; b; SM.B Me 18953, head with thick peduncle; c: SM.B Me 5601, head with partly degraded fibrous peduncle; d: SM.B Me 8469, head with thickened band near the top of the peduncle (arrow); e: SM.B Me 7050, pedunculate heads attached to a common axis; f: SM.B Me 21278, head with thickened band near the top of the peduncle (arrow); g: counterpart of the infructescence in f; h: SM.B Me 23879, portion of head fractured showing radial arrangement of compartments; i: counterpart of h, surface of head with 4–6 sided compartments; j–1: striate seeds; j: seed in situ from c, SM.B Me 5601; k: another seed from c; 1: detail from k, showing longitudinally elongate cells of surface seed coat giving striate appearance; scale bars: a–i = 1 cm; j–1 = 1 mm.

216


Plate 62 a–i: Carpolithus sp. 37, a–c: SM.B Me 4848; a: fruit showing smooth surface, with lower half partially broken revealing rounded locule and thick wall; b: detail of fruit wall and locule contents; c: detail of b, showing fruit wall composed of anticlinally aligned cells; d–i: SM.B Me 2299; d: fruit with base protruding into a short broad extension with large circular scar; e: reverse side of d; note rounded apex; f: fruit in d transversely sectioned; g: same, viewed from base, showing lensoidal outline; h: transverse section of fruit in f showing internal spongy viscous substance of uniform white color; i: detail of h. showing fruit wall composed of anticlinally arranged fibers whose ends form a fine polygonal pattern at the surface; scale bars: a, b, d–h = 1 cm, c, i = 1 mm.

218


Plate 63 a–h: Carpolithus sp. 38, a: SM.B Me 2092, globose fruit clothed by compact hairs; b: SM.B Me 11311, smaller specimen; c: SM.B Me 7261, fruit partially broken, showing full length of the enveloping hairs, and fracture through underlying yellowish white resinous layer; d: SM.B Me 15474, fruit with a prominent layer of yellowish white resin below the layer of splaying hairs; e: SM.B Me 15476, fruit partially broken exposing the layer of yellowish white resin; f: SM.B Me 15476, detail from e, showing rounded perianth lobes (arrows); g: SM.B Me 23336, fruit with resin visible as paler streaks through the fine hairs; h: SM.B Me 23336, detail from g showing three of the thin rounded tepals (arrows); scale bars: a–g = 5 mm; h = 3 mm.

220


Plate 64 a–m: Carpolithus sp. 39, a–c: SM.B Me 17448 a: fruit with at least two membranous valves or wings, with several seeds positioned near the center; b: detail of veins radiating from the central area of the seeds; c: detail of cluster of centrally placed seeds, and entire margin of the apical part of the membranous valve or wing; d: SM.B Me 4359, isolated circular seed with wrinkled surface; e: SM.B Me 4366, seed coat smooth, somewhat shiny. Specimen also used for SEM study; f: SM.B Me 2288, seed coat black, fractured and cracked due to brittle texture; g: SM.B Me 17448, detail of seed coat fragment from c, SEM showing outer layer of columnar sclereids with very thickened walls (no lumen visible) and inner layers of strongly compacted probably originally isodiametric fairly thick-walled cells, seed coat incomplete as not all layers were removed with the fragment; h: SM.B Me 4366, detail from e, seed coat with an outer columnar sclerotesta having small lumina near the inner edge, SEM; i: surface pattern from e of very fine polygonal isodiametric cell outlines, SEM; j: SM.B Me 17448, detail from c, seed surface layer with very fine polygonal pattern, and at least two circular openings, SEM. Compare with isolated seeds having the same surface pattern, e.g., k, m; k: detail from i, showing a small open pit which may be a stoma, hair base or site of former crystal, SEM; 1: SM.B Me 4359, enlargement from d, surface pattern of very fine polygonal isodiametric cell outlines; m: detail from 1, showing surface open pit comparable to j and k; scale bars: a–c = 1 cm; d–f = 5 mm; g = 200 µm; h = 80 µm; i = 100 µm; j, k, m = 20 µm; 1=50 µm.

222


Plate 65 a–c: Carpolithus sp. 40, SM.B Me 8956, a: ? infructescence showing opposite branching; b: detail of a, showing node with two lateral and one terminal bodies, right lateral body shows fibrous surface; c: detail of woody ? pedicel and multilayered surface of the left of the two lateral bodies; d–h: Carpolithus sp. 41, SM.B Me 2328 d: group of hard elliptical seeds within a berry-like fruit; e: detail of d, showing; a group of striated seeds; f: shiny ellipsoidal seed with longitudinally striate inner cuticular layer; g: another seed from same fruit; h: detail of g, showing wrinkly cuticle surface; scale bars: a–d = 1 cm; e = 2 mm; f, g = 1 mm; h = 0.25 mm.

224


Plate 66 a–p: Carpolithus sp. 42, a: SM.B Me 5740, rare example of fruit where black shiny layer shows very little blocky fracture; b: SM.B Me 21365, fruit fractured through the black shiny layer showing blocky fracture in section at the sides (arrows) and on the exposed fractured surface. Tissues of inner fruit wall protrude at apex and base; c: SM.B Me 21366, Fruit fractured through the black shiny layer showing blocky fracture at the right side. Brown tissues of inner fruit wall (paler colour) are revealed in the upper half of specimen where longitudinal fibre bands are also visible (arrows). Outermost fruit wall is present at the base (paler colour) where the specimen narrows to the attachment; d: SM.B Me 21350, fruit which became separated during preparation into an inner body (left of image, consisting of inner fruit wall) and an outer portion (consisting only of the black shiny blocky layer). A median fibre strand can be seen on the inner body (arrow) and is marked by a median groove on the outer portion; e: SM.B Me 4694, fruit showing longitudinal striations as a result of fractures in the black shiny blocky layer that is also seen in section at left of image. The outer fruit wall layer is present at the base (paler colour) where the specimen narrows to the former attachment; f: SM.B Me 21364, fruit showing blocky fracture of black shiny layer all over surface and with scattered overlying patches of outer fruit wall layer (paler colour); g: SM.B Me 21379, typical fruit lacking thin outer fruit wall and with clear blocky fracture of black shiny layer all over surface. No inner layers visible; h: SM.B Me 21327, digital oblique longitudinal section by SRXTM parallel with plane of compression. The outer fractured blocky layer can be recognised by the radial fractures (arrows). Most of the section passes through the underlying fruit wall layer which shows no cellular detail by SRTXM. The right half of the image has sectioned obliquely through the locule lining (paler colour more open tissue structure). The groove overlying the median fibre band is seen as a dark line; i: SM.B Me 5658, light microscopy, showing detail of transverse section of portion of fruit wall showing the diagnostic black shiny, apparently amorphous, layer with blocky fracture (base of image, arrow), underlying paler structured wall layer and distinctive innermost cellular locule lining of variable thickness. Photographed wet so locule closed as a result of compression; j: SM.B Me 21327, digital transverse section by SRXTM. The outer fractured black shiny blocky layer is scarcely distinguishable from the underlying layer by SRXTM in this plane of section (compare to h). The single locule is clearly seen as is the groove beneath the median fibre band. The locule lining is distinct (paler colour, more open tissue structure); k–o: SM.B Me 5658; k: SEM showing transverse section of fruit with distinctive fractured outer black shiny blocky layer (arrow), underlying wall layer and single locule now open after drying; l: SEM showing fine detail of black shiny blocky layer (from l) showing textured cell infill, possibly indicating secretory origin; m: detail from lower left portion of fruit wall from k showing black shiny blocky layer (base of image), underlying layer, the inner part of which shows fibrous nature, and strongly compact locule lining; n: detail of black shiny blocky layer from i showing barely recognisable cellular structure with cells completely infilled. (Detail of infill texture is shown in l); o: detail from lower right portion of fruit wall from k showing fruit wall layers, innermost cuticle with elongate cell pattern and locule lining tissues composed of platy cells; p: Sm.B Me 5690, SEM showing transverse section of fruit wall with black shiny blocky layer (base of image) underlying layer and clearly delimited locule. Locule lining not compacted, composed of equidimensional cells with thickened walls but open lumina; scale bars a–h = 5 mm; j = 3 mm; k = 500 µm; l = 1 µm; m = 20 µm; i, n–p = 200 µm.

226


Plate 67 a–d: Carpolithus sp. 43, SM.B Me 7060, a: very large elliptical fruit with multifragmented surface; b: detail of a, showing, elliptical seeds within interior of fruit; c: detail of the two smooth elliptical seeds from b; d: detail of cuticular surface of fruit; e–g: Carpolithus sp. 44, SM.B Me 16858, e: fusiform fruits borne on a twig with three leaves; f: detail of e, showing elongate fruit with longitudinal grooves, distally constricted and tapering above into an apical beak-like structure; g: detail of e, showing fruits that appear to be borne on the petiole, with an ellipsoidal body on the petiole (arrow); h–k: Carpolithus sp. 45, h: SM.B Me 20418, crescent shape fruit with truncate base and 3 visible valves separated by strong longitudinal ridges; i: detail of h, showing small 5-lobed pad of tissue (? stigma) at apex and ribbed surface; j: SM.B Me 25000; obovate fruit with truncate base and apex, and three visible valves with a median groove on the front facing valve; k: detail of valve from j showing smooth surface, median longitudinal groove and marginal ridge; scale bars: a = 3 cm; b, c, e, h–k = 1 cm; f, g = 5 mm; d = 0.5 mm.

228


Plate 68 a–d: Carpolithus sp. 46, SM.B Me 7073, a: large globose fruit, on a thick pedicel, with a hypogynous perianth of more than seven imbricate lobes; b: detail of a, showing a prominent longitudinal ridge with a median groove at the crest. Arrow indicates area of seeds shown in d; c: detail of a showing the overlapping perianth parts; d: seeds (arrows) and cuticular margin of fruit; e–h: Carpolithus sp. 47, SM.B Me 7302, e: ellipsoidal, bilaterally symmetrical specimen, rounded at one end, pointed at the other; f: removed from shale, showing ca. 18 rounded longitudinal ridges that extend to both ends; g: opposite side showing longitudinal ridges and groove; h: polar view showing D-shape; i: Carpolithus sp. 48, SM.B Me 23185, small fruit (possibly trigonal) with median longitudinal rib (?keel); scale bars: a–c = 1 cm; d = 1 mm; e–i = 3 mm.

230


Plate 69 a–f: Carpolithus sp. 49, a–c: SM.B Me 7621; a: infructescence bearing subsessile fruits; b: detail of a, showing obovoid fruits with persistent calyx, of which parts of three lobes can be seen in face view (arrows); c: detail of a showing fruit splitting longitudinally; d–f: SM.B Me 8950; d: infructescence bearing subsessile fruits; e: detail of d, showing fruit splitting longitudinally into multiple persistent valve like units; f: same, showing five valve like units (arrows) and faint cup-like structure corresponding to the perianth seen in b; g–h: Carpolithus sp. 50, SM.B Me 4791, g: widely ellipsoidal operculate seed; h: detail of g, showing operculum surrounded by a raised rim, and the centre of the operculum with a small columnar protrusion; scale bars: a, d = 1 cm; b, c, g = 5 mm; e = 3 mm; f = 2.5 mm; h = 2 mm.

232


Plate 70 a–f: Carpolithus sp. 51, a: SM.B Me 21397, two seeds on same specimen, upper one fragmentary, lower one more complete with subovoid outline; b: detail of lower seed from a, reoriented, with patches of thin leathery shiny outer cuticle; c: SM.B Me 21394, seed broken at one end but opposite end (base of image) not compressed; d: detail of b, showing patches of thin leathery shiny outer cuticle; e: detail of the upper seed from a, reoriented, and illuminated to show a thickened mound at one end (top of image); f: detail of c, showing mound with radiating columnar cells and a narrow central opening which expands inwards forming a vestibule; g–i: Carpolithus sp. 52, SM.B Me 21575, g: elliptical ribbed fruit with two lobes protruding from broken end; h: detail of lobes; i: detail of polygonal cellular pattern on surface; scale bars: a, g = 5 mm; b, c, = 3 mm; d–f = 2 mm; h = 1 mm; i = 0.5 mm.

234


Plate 71 a–f: Carpolithus sp. 53, a–c: SM.B Me 10523; a: SM.B Me 10523A, flattened globose fruit with fragments of radially aligned dark seeds; b: SM.B Me 10523B, counterpart of a; c: detail of b, showing clumps of flat radially aligned elliptical seeds; d: SM.B Me 7353, flattened globose fruit with remains of seeds; e, f: SM.B Me 7071; e: globose fruit with an elliptical seed remaining; f: enlargement of smooth elliptical seed from e which is apparently represented only by inner seed coat layer (outer layer missing); g–i: Carpolithus sp. 54, SM.B Me 23874, g: large cordate woody fruit with septum and two D-shaped locules, photographed in water; h: as g but photographed dry; i: detail of h, showing bulbous locule and fibrous wall; scale bars: a, b, d, e, g, h = 1 cm; c = 1 mm; f = 2 mm; i = 5 mm.

236


Plate 72 a, b: Carpolithus sp. 55, SM.B Me 25013, a: Four pyrenes or nutlets surrounded by a circular wing; b: detail of a, showing wrinkled surface of pyrenes and membranous wing; c–h: Carpolithus sp. 56, c: SM.B Me 4533, small round seed; d–f: SM.B Me 4138; d: orbicular seed; e: detail of d, showing elongate groove around the right-hand edge and finely pitted surface; f: detail of e, showing isodiametric poligonal cell outlines; g: SM.B Me 15115, cluster of four seeds; h: detail of g showing isodiametric polygonal cell outlines; scale bars: a = 1 cm; b = 5 mm; c, d, g = 3 mm, e = 1 mm; f = 200 µm; h = 0.3 mm.

238


Plate 73 a–o: Carpolithus sp. 57, a–e: seeds showing shiny surface and brittle fracture pattern; a: SM.B Me 4012, seed with slight faceting either side of a broad ridge (possible raphe ridge) and possible apical scar; b: SM.B Me 4031, seed showing brittle cracks but most fragments remain in original position in subglobose seed; c: SM.B Me 4354, badly broken specimen revealing seed coat layers; d: SM.B Me 4083, cracked specimen, flattened as a result of cracking; e: SM.B Me 5738, very rare example with small patches of surface striations; f: SM.B Me 4012, detail of a, showing very shiny surface with fine polygonal cell pattern; g, h: detail of e, with g showing area of striated surface and h showing detail of striations as undulating wrinkles and adjacent more typical surface shiny with fine polygonal cells (to right); i–o: SEM of seed surface and seed coat layers; i, j: SM.B Me 4346; i: seed coat surface with distinctive polygonal pattern; j: detail of i, showing polygons formed from rounded ridges enclosing a flatter area with 10 to 13 small evenly spaced pits, pits closed by outer periclinal wall of cell; k–n: SM.B Me 2631, details of inner seed coat layer which separates readily from outer layer; k: vertical fracture of main part of inner seed coat layer showing radially elongate fibres and outer surface of this layer with polygonal pattern; l: inner surface of inner seed coat layer, a thin cuticle with outlines of elongate cells; m: adjacent vertical aligned fibres and transverse sections through tangentially aligned fibres in main part of inner seed coat layer, both overlain by outermost part of inner seed coat composed of at least two layers of thin-walled, radially narrow, polygonal cells; n: detail of m showing transverse sections of fibres and overlying thin-walled cells; o: SM.B Me 4346, fractured vertical section of outer seed coat layer showing several layers of polygonal cells, outermost layer shows small lumina just beneath the outer periclinal wall of surface cell, each lumen linked to a small surface pit, these lumina might have housed crystals; scale bars: a–e = 2 mm; f = 250 µm; g = 1 mm; h = 0.5 mm; i = 20 µm; j = 10 µm; k = 100 µm; l = 40 µm; m = 200 µm; n = 40 µm; o = 50 µm.

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Plate 74 a–p: Carpolithus sp. 58, a: SM.B Me 2279, smooth shiny globose seed with sharp junction at base of apically mounded aril; b: SM.B Me 2281, arillate seed with broken seed coat; c: SM.B Me 5752, arillate seed broken and fragments displaced showing thin seed coat; d: SM.B Me 12, dry seed with prominent aril showing aligned cells radiating from attachment; e: SM.B Me 2280, specimen with patch of aril at left, a crucial link between fully arillate specimens and those specimens lacking the aril and just showing the centrally mucronate truncation; f: SM.B Me 7195, seed with a patch of aril preserved over the truncation; g: SM.B Me 7146, apical view of truncation, with central mound, aril missing; h: detail of aril from d showing radially aligned cell rows; i: SM.B Me 4731, obliquely compressed and partly fractured seed showing black shiny lateral surfaces, contrasting with dull texture of circular truncation; j: SM.B Me 4387, seed showing circular truncation with central protrusion; k: SM.B Me 11770, broken specimen with extended central protrusion; l: SM.B Me 5752, vertical fracture of seed coat from c showing cells aligned anticlinally; m–p: SM.B Me 5752, SEM; m: vertical fracture of seed coat showing thin smooth surface layer beneath which is very thick-walled tissue composed of multiple cells each with minute lumen; n: another vertical fracture of seed coat showing thick outer almost solid sclerotic layer, thinner central layer of more fragile, less sclerified tissue and innermost layer of compacted, thick-walled parenchymatous tissue; o: smooth surface with scarcely evident suggestions of possible polygonal cell pattern only visible when specimen carefully orientated at an angle in the SEM; p: innermost surface of inner seed coat layer, lined by a thin cuticle; scale bars: bar in d applies for a–g, bar in i applies also to j, = 5 mm; h, = 1 mm; l = 20 µm; m, n = 100 µm; p = 50 µm.

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Plate 75 a–d: Carpolithus sp. 59, a, c: SM.B Me 8803; a: pedicellate fruit borne on elongate twig; c: detail of a, showing upper half of fruit in face view with two meridional ribs and three apparent valves, and cup-like structure over the lower half of fruit; b, d: SM.B Me 2573; b: pedicellate fruit with associated elongate twig; d: detail of b, showing fruit with a prominent, coarsely longitudinally wrinkled cup-like structure preserved over the lower half; e, f: Carpolithus sp. 60, SM.B Me 8802, e: ellipsoidal fruit with densely verrucate surface, a patch of which is removed, revealing seed like structures; f: detail of e, showing closely packed barrel-shaped to ovoid seed-like structures; g: Carpolithus sp. 61, SM.B Me 8697, globose fruit on pedicel, with hypogynous disk; scale bars: a–g = 1 cm.

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Plate 76 a–d: Carpolithus sp. 62, SM.B Me 20618, a: fruit with two overlapping, elongate folded wings, with bulging fruit body at lower right; b: detail of fruit body from a, showing closely spaced longitudinal ribs on the fruit body and the attachment of wings near base of the left and right sides; c: detail of the wing surface from a, with subtle longitudinal striation and/or venation; d: detail of fruit body from a, showing longitudinal ribs at left and group of small spirally arranged overlapping rhomboidal scales in the upper central part between the wings; e, f: Equisetum bulbil, SM.B Me 4190, opposite ends of the same specimen, uncompressed structure would be unlikely in a fruit or seed which, due to lack of internal space for embryo, would tend to compress in oil-shales; e: shows circle of triangular lobes with central protrusion, which superficially resemble tiny calyx lobes; scale bars: a, b = 1 cm; c–f = 5 mm.

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Taxonomic index Ailanthus confucii Unger (Pl. 38l, m)..................................................50 Alangiaceae..........................................................................................17 Alangium sp. (Pl. 1d)...........................................................................17 Altingiaceae ........................................................................................17 Ampelopsis sp. (Pl. 43a–i)....................................................................54 Anacardiaceae .....................................................................................18 Anacardium germanicum Manchester, Wilde & Collinson (Pl. 2a–e).............................................................................................18 Annonaceae..........................................................................9, 10, 15, 64 Ant.........................................................................................................9 Aphananthe cf. tenuicostata Dorofeev (Pl. 7i–l)................................22 Apocynaceae........................................................................................20 Araceae......................................................................................6, 16, 70 Araliaceae...................................................................................... 11, 21 Arecaceae.............................................................................................20 Artiodactyla, artiodactyl, artiodactyls....................................................8 Aumelasia...............................................................................................8 bat............................................................................................................. Beckettia, cf. Reid & Chandler (Pl. 29h–l).........................................38 Berchemia mellerae sp. n. (Pl. 36h–n).................................................47 Betulaceae.......................................................................... 11, 18, 19, 69 Bignoniaceae........................................................................................21 bird...................................................................................................8, 13 Burseraceae..........................................................................................21 Bursericarpum sp. (Pl. 7g, h)...............................................................22 Buxaceae................................................................................................6 Buxolestes........................................................................................8, 13 Camelliacarpoidea messelensis sp. n. (Pl. 40a–i)................................50 Canarium sp. (Pl. 7a–f)........................................................................21 Cannabaceae........................................................................................22 Carpolithus callosaeoides (Engelhardt) comb. n. (Pl. 45e–l)............56 Carpolithus sp. 1..................................................................................57 Carpolithus sp. 2..................................................................................57 Carpolithus sp. 3..................................................................................58 Carpolithus sp. 4..................................................................................58 Carpolithus sp. 5..................................................................................58 Carpolithus sp. 6..................................................................................59 Carpolithus sp. 7..................................................................................59 Carpolithus sp. 8..................................................................................59 Carpolithus sp. 9..................................................................................60 Carpolithus sp. 10................................................................................60 Carpolithus sp. 11................................................................................60 Carpolithus sp. 12................................................................................60 Carpolithus sp. 13................................................................................61 Carpolithus sp. 14................................................................................61 Carpolithus sp. 15................................................................................61 Carpolithus sp. 16................................................................................61 Carpolithus sp. 17................................................................................62 Carpolithus sp. 18................................................................................62 Carpolithus sp. 19................................................................................62 Carpolithus sp. 20................................................................................63 Carpolithus sp. 21................................................................................63 Carpolithus sp. 22................................................................................63 Carpolithus sp. 23................................................................................64 Carpolithus sp. 24................................................................................64 Carpolithus sp. 25................................................................................64 Carpolithus sp. 26................................................................................64 Carpolithus sp. 27................................................................................64 Carpolithus sp. 28................................................................................65 Carpolithus sp. 29................................................................................65 Carpolithus sp. 30................................................................................65 Carpolithus sp. 31................................................................................65 Carpolithus sp. 32................................................................................66 Carpolithus sp. 33................................................................................66 Carpolithus sp. 34................................................................................67 Carpolithus sp. 35................................................................................67

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Carpolithus sp. 36................................................................................67 Carpolithus sp. 37................................................................................68 Carpolithus sp. 38................................................................................68 Carpolithus sp. 39................................................................................68 Carpolithus sp. 40................................................................................69 Carpolithus sp. 41................................................................................69 Carpolithus sp. 42................................................................................70 Carpolithus sp. 43................................................................................70 Carpolithus sp. 44................................................................................70 Carpolithus sp. 45................................................................................71 Carpolithus sp. 46................................................................................71 Carpolithus sp. 47................................................................................71 Carpolithus sp. 48................................................................................71 Carpolithus sp. 49................................................................................72 Carpolithus sp. 50................................................................................72 Carpolithus sp. 51................................................................................72 Carpolithus sp. 52................................................................................72 Carpolithus sp. 53................................................................................73 Carpolithus sp. 54................................................................................73 Carpolithus sp. 55................................................................................73 Carpolithus sp. 56................................................................................73 Carpolithus sp. 57................................................................................74 Carpolithus sp. 58................................................................................74 Carpolithus sp. 59................................................................................75 Carpolithus sp. 60................................................................................75 Carpolithus sp. 61................................................................................76 Carpolithus sp. 62................................................................................76 Cayratia jungii (Gregor) Chen & Manchester (Pl. 43o, p)................54 Cedrelospermum leptospermum (Ettingshausen) Manchester emend. Wilde & Manchester (Pl. 41a–f)....................................................52 Cleyera sp. (Pl. 36e–g).........................................................................47 Cocculus lottii sp. n. (Pl. 32k–r)..........................................................43 colliiform................................................................................................9 Conifers......................................................................................7, 18, 19 coraciiform............................................................................................... Corylopsis maii sp. n. (Pl. 14a–e)........................................................25 Corylopsis waltheri sp. n. (Pl. 14f–t)...................................................26 Crassivitisemen wildei (Chen & Manchester) comb. n. (Pl. 44a–m, 45a–d)..............................................................................................55 Cruciptera schaarschmidtii Manchester, Collinson & Goth (Pl. 20d–g, 21b)......................................................................................32 Cyclanthaceae......................................................................................22 Cyclanthus messelensis S.Y. Smith, Collinson & Rudall (Pl. 8a–e)........22 Cyperaceae...........................................................................................23 Cypselites sp. (Pl. 4a–e).......................................................................20 Darmstadtia biseriata gen. et sp. n. (Pl. 6a–c, e).................................21 Darwinius...............................................................................................8 Decodon, cf. (Pl. 25a–f, 26a–g)...........................................................35 Diploclisia rugulosa sp. n. (Pl. 31a–d)................................................39 Doliostrobaceae ..................................................................................21 Doliostrobus taxiformis (Sternberg) Z. Kvaček emend. Kunzmann (Pl. 1a–c)..........................................................................................21 Elaeocarpaceae.....................................................................................23 Eocoracias.............................................................................................8 Eoglaucidium.........................................................................................9 Eogliravus..............................................................................................8 Eomastixia cf. rugosa Chandler emend. Chandler (Pl. 28a–m 29a–g).....36 Equisetum sp. (Pl. 75e, f).....................................................................76 Ericaceae....................................................................................7, 12, 72 Euphorbiaceae......................................................................................24 Euphorbiotheca gothii sp. n. (Pl. 10a–l, 11a–p)..................................24 Eurodexis...............................................................................................8 Eurohippus.............................................................................................8 Eurotamandua......................................................................................20 Fagaceae.....................................................................................7, 12, 71 Fern.......................................................................................................... Flacourtiaceae.................................................................... 11, 15, 59, 61 Friedemannia messelensis gen. et sp. n. (Pl. 5a–n).............................20

gruiform.................................................................................................8 Hamamelidaceae..................................................................................26 Hooleya sp. (Pl. 21a)............................................................................32 Icacinaceae . ........................................................................................27 Icacinicarya densipunctata sp. n. (Pl. 19f–k)......................................31 Icacinicarya sp. (Pl. 20a–c).................................................................31 Icacinicarya tiffneyi sp. n. (Pl. 17a–i)..................................................29 Incertae Sedis.......................................................................................56 insect........................................................ 8, 9, 11, 17, 32, 55, 57, 58, 71 Juglandaceae........................................................................................32 Karinschmidtia rotulae gen. et sp. n. (Pl. 33a–o)................................42 Kopidodon..............................................................................................8 Lannea hessenensis sp. n. (Pl. 2g–i)....................................................19 Lauraceae.............................................................................................37 Lauraceae genus indet. 1 (Pl. 21c–d)...................................................34 Laurocarpum sp. 1. (Pl. 22a–m)..........................................................33 Laurocarpum sp. 2. (Pl. 22n)...............................................................33 Laurocarpum sp. 3. (Pl. 21e–g)...........................................................33 Leguminocarpon herendeenii n. sp. (Pl. 24a–d)..................................34 Leguminocarpon sp. 1 (Pl. 24e, f)........................................................35 Leguminocarpon sp. 2 (Pl. 24g)...........................................................35 Leguminosae .......................................................................................38 Lythraceae............................................................................................39 Magnolia sp. 1 fruit (Pl. 27g)...............................................................36 Magnolia spp. seeds (Pl. 27a–f)...........................................................36 Magnoliaceae.......................................................................................36 mammal................................................................................8, 13, 15, 16 mantid fly................................................................................................. Martinmuellera tuberculata gen. et sp. n. (Pl. 31j–m)........................40 Masillamys...................................................................................6, 8, 16 Mastixiaceae........................................................................................36 ?Mastixia sp. (Pl. 27h–t)......................................................................37 mastixioid indet. (pls 29n, o; 30j, n)....................................................38 Mastixiopsis nyssoides Kirchheimer (Pl. 30a–i)..................................38 Meliosma sp. (Pl. 38a–f)......................................................................49 Menispermaceae .................................................................................38 Menispermeae indet. (pl. 31i)..............................................................40 Menispermaceae, unnamed (Pl. 34k)...................................................44 Messelobunodon..................................................................................12 Messelornis..........................................................................................12 Mimosites spiegeli Engelhardt (Pl. 23a–f).........................................34 Myricaceae....................................................................................... 6, 11 Myristicacarpum sp. (Pl. 34o, p).........................................................46 Myristicaceae.......................................................................................46 Mytilaria boglei sp. n. (Pl. 12, 13a–f)..................................................26 Natsiatum, cf. (Pl. 16k–m)...................................................................28 Nymphaeales, Unnamed seeds (Pl. 35a–i)...........................................46 Nyssa disseminata (R. Ludw.) Kirchheimer (Pl. 36a–d).....................47 Nyssaceae.............................................................................................47 Palaeocarya sp. sensu Manchester et al. (Pl. 20h, i)..........................33 Palaeohosiea bilinica (Ettingshausen) Kvaček & Bůžek (Pl. 15a–k, m, n, 16a–c).....................................................................................27 Palaeohosiea sp. (Pl. 16d–j)................................................................28 Palaeophytocrene sp. (Pl. 17j–p).........................................................29 Palaeosinomenium ornamentum sp. n. (Pl. 32b–d).............................41 Palaeosinomenium sp. (Pl. 32f–i)........................................................41 Palaeosinomenium venablesii Chandler (Pl. 32e)..............................41 Palaeovitis sp. (Pl. 43j–n)....................................................................54


pantolestid..............................................................................................8 Parabaena cf. europaea Czeczott & Skirgiello (Pl. 34d–f)..............44 paroxoyclaenid.......................................................................................8 Parthenocissus britannica (Heer) Chandler (Pl. 42i–k)....................53 Perissodactlya, perissodactyl, perissodactyls.......................................12 ?Pericampylus sp. (Pl. 32j)..................................................................42 Pentaphylacaceae ................................................................................51 Pentoperculum minimus (Reid & Chandler) Manchester (Pl. 3a–h).......18 Phytocrene punctilinearis sp. n. (Pl. 18a, b, d, e, h–j).........................30 piciform..................................................................................................8 Pinaceae...........................................................................................7, 12 Platanaceae.................................................................................7, 14, 15 Pleiogynium mitchellii sp. n. (Pl. 3i–m)..............................................19 Primate...................................................................................................8 Primobucco............................................................................................8 Primozygodactylus.................................................................................9 Propalaeotherium..................................................................................8 Pyrenacantha sp. 1 (Pl. 18c, f, g).........................................................30 Pyrenacantha sp. 2 (Pl. 18k, l; 19a–d).................................................30 Pyrenacantha, cf. (Pl. 19e)..................................................................31 Restionaceae........................................................................................16 Rhamnaceae.........................................................................................47 rodent.................................................................................12, 13, 20, 78 Rutaceae...............................................................................................48 Rutaceae undetermined A sensu Collinson & Gregor (Pl. 37f).........49 Rutaceae undetermined B sensu Collinson & Gregor (Pl. 37g–i).....49 Rutaspermum chandleri Collinson & Gregor (Pl. 37m)....................48 Rutaspermum messelense Collinson & Gregor (Pl. 37a–e)...............48 Sabiaceae ............................................................................................53 sandcoleid..............................................................................................9 Sapindaceae..................................................................19, 65, 66, 73, 79 Sapotispermum, cf. (Pl. 38j, k)............................................................49 Saportaspermum sp. (Pl. 46a–h)..........................................................56 Sapotaceae............................................................................................49 Simaroubaceae ....................................................................................50 Sloanea messelensis sp. n. (Pl. 9a–o). ................................................23 Spirellea sp. (Pl. 46i–m)......................................................................57 Steinhauera subglobosa Presl. emend. Mai (Pl. 1e, f)........................17 Stephania hootae sp. n. (Pl. 31e–h).....................................................39 Symplocaceae........................................................................................7 Tapiscia pusilla (Reid & Chandler) Mai (Pl. 39a–l)...........................50 Tapisciaceae.........................................................................................50 Tetraedron........................................................................................7, 12 Theaceae..............................................................................................50 Tinomiscoidea jacquesii sp. n. (Pl. 34a–c)...........................................44 Tinosporeae sp. 1 (Pl. 34g, h)..............................................................45 Tinosporeae sp. 2 (Pl. 34i, j)................................................................45 Tinosporeae sp. 3 (Pl. 34l–n).............................................................445 Toddalia, cf. (Pl. 37j–l)........................................................................49 Toricellia bonesii (Manchester) Manchester (Pl. 39m–p) . ..............51 Toricelliaceae.......................................................................................51 Ulmaceae..............................................................................................52 Vitaceae................................................................................................52 Vitaceae, unnamed (Pl. 43q, r).............................................................55 Vitis messelensis sp. n. (Pl. 42a–h). ....................................................52 Volkeria messelense Smith, Collinson, Simpson, Rudall, Marone & Stampanoni (Pl. 8f–i).......................................................................23 Wardensheppeya sp. (Pl. 32a)..............................................................42 weevil....................................................................................................... Zygodactylidae.......................................................................................9

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