Vegetative and reproductive structure of the extinct Platanus neptuni ...

Plant Syst. Evol. 244: 1–29 (2004) DOI 10.1007/s00606-003-0082-2

Vegetative and reproductive structure of the extinct Platanus neptuni from the Tertiary of Europe and relationships within the Platanaceae Z. Kvacˇek1 and S. R. Manchester2 1 2

Charles University, Praha, Czech Republic Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA

Received June 2, 2002; accepted October 6, 2003 Published online: January 28, 2004 Springer-Verlag 2004

Abstract. Platanus neptuni (Ettingshausen) Bu˚zˇek, Holy´ & Z. Kvacˇek was a conspicuous warmtemperate to subtropical element of Late Eocene to Late Miocene European floras. In our concept, the P. neptuni plant includes not only globose infructescences upon which the species epithet is based, but also staminate and pistillate inflorescences and distinctive stipulate foliage. The leaves range from simple (P. neptuni morphoforma reussii (Ett.) comb. et stat. nov.) to trifoliolate (P. neptuni morphoforma fraxinifolia (Johnson & Gilmore) comb. et stat. nov.) and sometimes quinquefoliolate (P. neptuni morphoforma hibernica (Johnson & Gilmore) comb. et stat. nov.) with unlobed elliptical to obovate laminae that are uniform in venation, marginal serration, and epidermal structure. Foliar twigs confirm that the leaves are deciduous, with each petiole base enveloping a bud, as in extant Platanus subgen. Platanus. Platanus neptuni differs from extant species of the genus by large peltate glandular trichomes on the fruits and leaves, a prominent circumscissile rim on the stalk below the inflorescence, as well as by the tendency for compound foliage. These characters justify its position within an extinct subgenus of the Platanaceae (Platanus L. subgen. Glandulosa Z. Kvacˇek, Manchester & Guo). Platanus neptuni was common in mesic humid subtropical forests on volcanogenic subtrates and at sea shores.

Key words: Platanaceae, mid-Tertiary, foliage, reproductive organs, morphology, anatomy, palaeoecology.

The family Platanaceae has an excellent fossil record extending to the early Cretaceous based on leaves and reproductive parts and it is clear that the family was more diverse and widespread in the past than it is today. Even in the Tertiary, the family included taxa that are extinct, including forms with compound leaves like Platanites Forbes (Crane et al. 1988, McIver and Basinger 1993), and Platanus bella (Kvacˇek et al. 2001). The Platanus neptuni complex seems to show transition between simple and compound foliage within the same lineage. This unusual platanaceous species was first recognized from the Oligocene and Miocene of northern Bohemia by Bu˚zˇek et al. (1967). These authors noted that elliptical to obovate, simple leaves could be linked by similarities of epidermal structure with cooccurring inflorescences and infructescences called Sparganium neptuni by Ettingshausen (1866) and demonstrated that the staminate inflorescences contained abundant pollen matching that of extant Platanus. Establishing the new combination, Platanus neptuni

Z. Kvacˇek and S. R. Manchester: Platanus neptuni complex from the Tertiary of Europe

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(Ettingshausen) Bu˚zˇek, Holy´ & Z. Kvacˇek, the same authors noted that although the fossil species differs from most modern species of Platanus, which possess lobed leaves, similar unlobed leaves occur in the extant species P. kerrii from Laos and Viet Nam. Subsequently, the leaves, inflorescences and infructescences of P. neptuni have been reported from numerous additional localities extending the palaeogeographic range in Germany, Ireland, Denmark, Hungary, Austria, Romania, Bulgaria and France (Fig. 1, Table 1 and Appendix). As more specimens of Platanus neptuni and related taxa have been investigated, we now realize that this taxon intergrades with similar early Paleocene species of Platanus that had compound leaves (Kvacˇek et al. 2001). From the Tertiary of western and central Europe, Walther (1985) recognized compound leaves of three to five leaflets with the same epidermal anatomy as P. neptuni, as an independent species, P. fraxinifolia. Platanus bella (Heer) Z. Kvacˇek, Manchester & Guo (2001) was a similar species with trifoliolate leaves from the Paleocene of North America, Greenland, and Asia. Kvacˇek et al. (2001) erected the new subgenus, Glandulosa Z. Kvacˇek, Manchester & Guo to accommodate -5˚

0˚

5˚

10˚

P. bella, P. fraxinifolia, and P. neptuni because of shared epidermal characters in these taxa that set them apart from the two extant subgenera. Although reproductive organs remain unknown for P. bella, nicely preserved inflorescences and infructescences are associated with P. neptuni, which now provide the opportunity for a more informed assessment of the affinities of P. subgen. Glandulosa within the Platanaceae. The purpose of this paper is to summarize all information on the morphology and epidermal anatomy of the Platanus neptuni complex, and to describe and illustrate the vegetative and reproductive structure more fully for comparison with other fossil and extant members of the Platanaceae. We propose a refinement of the nomenclature for this composite plant, recognizing P. neptuni as a single species that included three leaf forms named: morphoforma reussii (simple leaves), morphoforma fraxinifolia (trifoliolate leaves) and morphoforma hibernica (quinquefoliolate leaves). We review its distribution in time and space based on reliable fossil records in Europe, augmented by reference to a newly recognized related population in North America. Finally, we add 15˚

20˚

25˚

5 55˚ 1 6 3

50˚

11 10

4

7 8 9 14 13 12 18 17

45˚ 2

15 16 21 22 23 20 24

19

25 26 27

km 40˚

0

29 500

28 30 31

Fig. 1. Distribution map of the Platanus neptuni complex in Europe. The numbers correspond to localities or groups of localities indicated in Table 1 (for detailed locations see Hably 1985c, Mai 1995, Givulescu 1997 and original sources in appendix)


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Table 1. Review of European localities containing Platanus neptuni, indicating geologic age and the kinds of organs recovered. (numbers in brackets [] refer to locations on the map in Fig. 1 and references in Appendix) Locality

Age

Vegetative organs

Kucˇlı´ n (type locality) [9a]

Late Eocene K-Ar Leafy shoots, simple age > 38.3 ± 0.9 and trifoliolate (rare) my leaves, stipules Hlinna´, core Uc 9 [9b] L Eocene Simple leaves Klausa [7a] L Eocene Quinque- and trifoliolate leaves Bo¨hlen, Borna, Borna-Ost, L Eocene Zeitz Simple leaves Witznitz [7b] ‘‘Floren-komplex’’ Stare´ Sedlo, Cˇesky´ Chloumek, Novy´ Kostel [8a]

L Eocene

Flo¨rsheim [3]

E Oligocene NP 23 zone

Haselbach, Schleenhain, Beucha [7c]

E Oligocene Haselbach ‘‘Floren-komplex’’ E Oligocene Simple leaves NP 24 E Oligocene Simple leaf Simple leaves E Oligocene K-Ar age > 32.7 ± 0.8 my

Bystrˇ ice/Olsˇ ı´ [15] Bechlejovice [9c] Kundratice [9d]

Present Present Present

Absent

Present Absent Present

Simple leaves

Absent Staminate ?, pistillate inflorescences, infructescences, achenes Staminate, pistillate inflorescences, achenes, stamens Pistillate inflorescences, infructescences, isolated achenes Infructescences

Simple leaves, stipules

Absent

Absent

Simple leaves, stipule Simple leaves Simple leaves

Infructescence Absent Absent

Absent Absent Absent

Simple leaves, ? leaflets, stipules

Seifhennersdorf [10b]

E Oligocene K-Ar age > 30.4 ± 1.5 my E Oligocene K-Ar age > 29.5 ± 1.5 my Oligocene K-Ar age 29–30 MA Oligocene L Oligocene E Oligocene MP 21

Simple leaves

Stadice, Lochocˇice [9f] Zˇichov [9g] Dve´rce [8b]

Staminate inflorescences, infructescences, isolated achenes Absent

Absent

Present

Oligocene

Suletice, Suletice-Berand, Holy´ Kluk hill [9e]

Pistillate inflorescences, infructescences Absent Absent

Simple, trifoliolate (rare), quinquefoliolate (rare) leaves & dispersed cuticles Simple and Absent trifoliolate (rare) leaves Fragmented simple Absent leaves

Markvartice [10a]

Knı´ zˇecı´ -Pirskenberg hill [10c]

Reproductive organs Cuticle

Present

Present

Present

Present

Absent

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Table 1 (continued) Locality

Age

Vegetative organs


Bois d’Asson [2]

E L Oligocene boundary L Eocene & E Oligocene E Oligocene

Simple leaves

Absent

Absent

Simple (rare) and trifoliolate leaves Simple leaves

Absent

Absent

Absent

Absent

Simple and trifoliolate leaves Simple and trifoliolate leaves Simple leaves

Absent

Absent

Absent

Absent

Absent

Absent

Simple leaves

Absent

Present

Simple leaves Simple leaf Simple, trifoliolate quinquefoliolate (rare) leaves Simple leaf Fragmented simple leaves, a trifoliolate leaf Absent


Absent Absent Present

Infructescence Infructescences

Absent Present

Infructescences, dispersed fruitlets

Absent

Simple leaves Simple leaves

Absent Absent

Present Present

Simple leaves

Absent

Absent

Trifoliolate, ? simple leaves Simple leaves

Absent

Absent

Absent

Absent

Simple leaves

Absent

Absent

Simple and trifoliolate leaves Simple leaves

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Mesta Graben-Eleshnitsa [29a] Mesta Graben- Boukovo [29b] Polkovnik Serafimovo Graben [30a] Hvojna Basin-Pavelsko [30b] Socka [18]

E Oligocene

L Eocene to E Oligocene L Eocene to E Oligocene Budapest [20] E Oligocene NP 23 Eger-Kiseged hill [23a] E Oligocene NP 23 Noszvaj Hercegoldal [23b] E Oligocene Lough Neagh, core, L Oligocene Washing Bay [1] (palyno-dating W. Krutzsch) Rott [4] L Oligocene Bockwitz, Borna-Ost, L Oligocene Delitzsch-Su¨d, Thierbach Espenhain-Sto¨rmhal [7e] ’’Floren-komplex’’ Cores Spremberg 37/60, L Oligocene Jahmen NSL 42/65, Nochten W [11a] Core Kleinsaubernitz [11b] L Oligocene Linz, Eferding [13] L Oligocene NP 25 – NN1 Krumvı´ rˇ [14] L Oligocene Egerian Brezno [16a] L Oligocene Eger-Wind clay pit [23c] Andornakta´lya [23d] Vero¨cemaros [21a] Keszto¨lc [21b] Nagysa´p [21c] Ve´rtesszo¨lo¨s [21d] Poma´z [21e] Savine [17]

L Oligocene NP 24 – NP 25 L Oligocene Egerian L Oligocene Egerian L Oligocene Egerian L Oligocene Egerian L Oligocene Egerian L Oligocene NP 25 Oligocene

Simple and trifoliolate leaves Simple leaves Simple and trifoliolate leaves Simple leaf


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Table 1 (continued) Locality

Age

Vegetative organs


Divljana [27] Janda-Frushka Gora [25]

Oligocene L Oligocene

Absent Absent

Absent Absent

Evros volcanic complex [31] Witznitz, Laussig, Delitzsch-Su¨d, Goitsche, Muldenstein [6, 7d] Cores Spremberg 9/58, NSL 13/63 Daubau [11b] Bı´ lina Mine [9h]

L Oligocene

Simple leaves Simple, trifoliolate and pentafoliolate (rare) leaves Simple leaf

Absent

Absent

L Oligocene to E Miocene

Leaves simple, trifoliolate (rare)

Infructescences

Present

E Miocene

Absent

Infructescences

Absent

E Miocene

Absent

Absent

Dobrˇ ı´ cˇ at Plzenˇ [8e] Plesna´, core V 146 [8c]

E Miocene (?) E Miocene (?)

Simple and trifoliolate (rare) leaves Leaf fragment Leaf fragments

Present Present

Habartov, Dukla Mine, Pochlovice, Cheb Basin cores HV 1–7, 9, 11, 13, 14 [8d] Velka´ Cˇausa [16b]

E Miocene MN 4-5

Simple leaves

Absent Pistillate and staminate inflorescences Infructescences

E Miocene Eggenburgian E Miocene E Miocene M Miocene

Simple leaves

Absent

Absent

Simple leaves Simple leaves, twigs Absent

Absent Absent Present

Simple leaves Simple leaves Simple leaves

Absent Absent Staminate inflorescence, infructescence, achenes Absent Absent Absent


Dispersed cuticles

Absent

Present

Leaf fragments, dispersed cuticles Simple leaves

Absent

Present

Absent

Present

Lipovany [21a] Ipolytarnoć [21b] Fasterholt, Søby [5]

Lintsching [12] Sused [19] Beograd, Negotin [26] Ruzˇinci [28] Delureni, Valea de Cris [24a] Cornit¸ el [24b]

E-M Miocene M Miocene M Miocene Sarmatian M Miocene Sarmatian L Miocene L Miocene

our interpretation of its ecological amplitude based on associated vegetation and sedimentary environments. Material and methods Fossil remains of Platanus neptuni are known from numerous localities in the Tertiary of Europe (see

Present

Table 1 and Appendix). For this investigation, we focused on the specimens from Kucˇlı´ n, the type locality of P. neptuni, as well as specimens from selected localities providing the best preserved examples of both leaves and reproductive material. The Kucˇlı´ n diatomite is considered Late Eocene, based on radiometric dating of ca. 38 my (Bellon et al. 1998) and, together with the nearby core Uc 9

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at Hlinna´, is assigned to Late Eocene also on the basis of the occurrence of Doliostrobus and palynological data (Kvacˇek 2002). Other Czech localities studied include Oligocene sites, e.g. Bechlejovice (Knobloch 1994), Kundratice (Kvacˇek and Walther 1998), Markvartice (Bu˚zˇek et al. 1976), Suletice (Kvacˇek and Walther 1995) and some others more recently excavated, such as the Early Miocene Most Formation at the Bı´ lina Mine, Most Basin (Kvacˇek 1998), the Cypris Claystone of the Cheb and Sokolov basins (Bu˚zˇek et al. 1967, 1996) and the Plesna´ borehole V 146 in the Cheb basin periphery (Bu˚zˇek et al. 1982). Localities studied from Germany include Klausa and other sites in Saxony (Late Eocene to Early Miocene – Mai and Walther 1985, 1991; Mai 1997, 1999) and Flo¨rsheim in the Mainz Basin (Early Oligocene – Engelhardt 1911; Kvacˇek in press). Material studied from Hungary included collections from the Early Oligocene Tard Clay of Budapest, Late Oligocene (Egerian) sites, and the Early Miocene tuffite of Ipolytarnoć (Hably 1982, 1985a, 1990, etc.). A new record is available from the Late Oligocene volcanic complex of Evros, northeastern Greece (Kvacˇek and Velitzelos in progress, material stored at Athens University). We inspected also one of the type specimens of Dewalquea hibernica from Washing Bay (Hunterian Museum, Glasgow – GLAHM) for which new preparations of cuticles were obtained. Several type and voucher specimens from other Tertiary sites of the former Austrian monarchy (Socka, Tamsweg) were examined in the Joanneum Museum, Graz and Natural History Museum, Vienna. The specimens studied from Bohemia are housed in Prague at the Faculty of Science, Charles University (PRC), National Museum (NM), Czech Geological Survey (UUG), District Museum, Litomeˇrˇ ice (ML), Headquarters of the Bı´ lina Mines (DB), the private collection of Mr. J. Valı´ cˇek, Most, as well as the Natural History Museum, Vienna (NMW) and Geological Survey, Vienna (GBW). Specimens examined from the Hungarian sites are housed in the Hungarian Natural History Museum, Budapest (PB) and Ma´tra Museum, Gyo¨ngyo¨s. Those from German localities were examined in the Forschungsinstitut Senckenberg, Frankfurt/M. (SM. B), Geological Institute, Cologne University (GIK), Museum fu¨r Naturkunde der Humboldt-Universita¨t, Berlin (MfN) and Staatliche Naturhistorische Sammlungen, Museum fu¨r Mineralogie und Geologie, Dresden (MMG).

Impression and compression specimens were studied as exposed in cleaved shale, with some additional preparation to expose the margins of leaves and the styles and anthers of inflorescences using a fine needle under the dissecting microscope. Most microscopic slide preparations were available from the previous studies on the material from core Uc 9 at Hlinna´, Markvartice, Kundratice, Plesna´ core V 146, the Cypris Claystone in Czech Republic and Flo¨rsheim in Germany. Additional macerations were made on fructifications to reveal the pubescence, and on male inflorescences to obtain pollen in situ for scanning electron microscopy (SEM). To prepare cuticles for observation of epidermal structure, fragments of the fossils were removed and cleaned in HF, transferred into Schulze solution for a short time, rinsed in water, cleared in KOH solution, rinsed again in water, and mounted either on a glass microscope slide in glycerine with the coverslip framed by Canada balsam for transmitted light microscopy, or on aluminum stubs for SEM. Similarly, parts of staminate compression fossils were removed, cleaned in HF, and macerated with NaOH to reveal clumps of pollen which were mounted on glass slides for light microscopy or transferred to aluminum stubs for SEM. To reduce mechanical damage, the whole maceration process was done right on the microscope slide in drops of individual reagents (see Kvacˇek in press).

Results Systematic descriptions Reproductive material (Figs. 2–4, 8a). Both staminate and pistillate inflorescences are single, stalked, globose to ellipsoidal heads with a swollen rim at the junction of stalk with head. Epidermis of the stalk of both staminate and pistillate inflorescences (Fig. 8a) thickly cutinized, finely granulate on the surface. Epidermal cells of the stalks ± isodiametric quadrangular, rarely polygonal, ca. 20–30 lm across, arranged in longitudinal rows, anticlinal walls deeply cutinized, seen as shallowly undulate double lines. Trichome bases ± densely spaced, compound, consisting of isodiametric longitudinally oriented pairs or groups of smaller basal cells and showing the


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Fig. 2. Stalked pistillate inflorescences and infructescences of Platanus neptuni. a Orig. Engelhardt 1885, pl. 2, Fig. 2, showing protruding styles. Kundratice, ML 124. b Infructescence with differential development of the fruits. Dukla Mine, Habartov, NM G 7090a. c Infructescence with long stalk and protruding styles. Klausa, MMG KL 359:16. d Infructescence showing remnants of perianth and achenes arising from receptacular core with receptacle scars suggesting three or four achenes per flower, Kundratice, MMG KU 490a. e Enlargement from a, showing prominent circumscissile rim at junction of stalk and head (arrow). f Pistillate inflorescence showing protruding styles [note unusually loose arrangement of achenes]. Kundratice, PRC CKU 150. g Infructescence showing styles arranged in small groups in the lower left periphery, probably corresponding to each flower. Kucˇlı´ n, lectotype BP 55.2491.1. h Infructescence with most of the achenes shed from an ellipsoidal receptacular core, and pronounced rim. Kucˇlı´ n, DB 29.4.95. i Possibly abortive inflorescence with ellipsoidal head and pronounced rim below (arrow). Seifhennersdorf, MMG A2/9/17. Scale bars ¼ 5 mm

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b Fig. 3. Infructescences and achenes of Platanus neptuni. a Relatively mature infructescence nearing maximum size. Kundratice, NM G 7915a. b Remains of an infructescence from which the achenes have been shed, retaining prominent perianth. Note tripartite pattern on the bases of individual flowers. Kucˇlı´ n, syntype, BP 59.1194.1. c Three-dimensionally preserved specimen showing well developed perianth (fused tepals) surrounding each flower. Plesna´ V 146/44–45 m, NM G 8104. d Enlargement from c, showing smooth perianth surface and remnants of apparently three styles per flower (arrows). e Dispersed achenes with ellipsoidal body and elongate persistent style. Note fine trichomes arising from basal portion (arrows). Markvartice, NM G 8101. f Sieved specimen showing obovate outline. Borna-Ost, MMG 8172A. g Isolated achene with a coalified seed in lower half of ovary. Kundratice, PRC CKU 196. h Isolated achene retaining cuticle structure (shown in Fig. 3k). Markvartice, UUG MR 1242a. i Detail of surface of an achene showing peltate glandular trichomes and sparsely distributed trichome bases. Borna-Ost, MMG 8172A. j Enlargement from Fig. 3f, showing dense pubescence and peltate glandular trichomes on the lower portion of achene. k Exocarp cuticle from achene in Fig. 3h showing fragmentary trichome bases. Markvartice, PRC MR1242a/2. Scale bar in a ¼ 10 mm, b ¼ 5 mm, c, e–h ¼ 2 mm, d, i, j ¼ 1 mm, k ¼ 100 lm

rounded trace of simple stalk, ca. 25 lm across, without terminal trichome head preserved. Stomata widely scattered, longitudinally arranged, c. 50 lm long, squashed from sides, with long aperture (Fig. 8a), recalling the stomata on foliage and stipules. The cuticle in the place above the rim thinner than elsewhere. Pistillate inflorescences – infructescences (Figs. 2–3). Heads 20 mm diameter at max., stalk 23 mm long at max., flowers with perianth well developed, styles of individual carpels extending well beyond perianth. Number of pistils per flower difficult to observe directly in most specimens; clearly three in some specimens, including those from the type locality of Kucˇlı´ n (Fig. 3b) and from the Plesna site (Fig. 3d), yet up to 8 in the Fasterholt specimens (see discussion). Styles protruding, curved (Fig. 2f). Receptacular cores ellipsoidal, 2.5–6.5 mm wide and 3.5– 7.5 mm long (Fig. 2h). Abortive pistillate inflorescences common, typically with thick, short, truncate styles (Fig. 2i), and sometimes with well-developed perianth (Fig. 3b). Fruits spindle-shaped, 4–7 mm long, with variably positioned maximum width and long tapering slightly curved style, single-seeded (Fig. 3g), covered partly with peltate glandular and nonglandular trichomes, arising sparsely over the basal 1/3 (Fig. 3h–k) and consisting of a uniseriate base and thinner, incompletely preserved terminal part (Fig. 3k). The upper part

of the achene smooth, thinly cutinized, consisting of straight-walled narrow elongate cells about 20 lm wide and variously long with acute ends. Staminate inflorescences (Fig. 4). Heads ± globose (Fig. 4a–g), 46 mm at max., stalk 8–12 mm long; inflorescence composed of flowers which together bear numerous stamens. Perianth mostly poorly developed, short, not affecting the arrangement of stamens within the head, more prominent in immature heads. Stamens observed both within the inflorescence and isolated in the sediment, with very short filaments, thin-walled longitudinal lateral pollen sacs and domed to conical connectives. Epidermis of the connective moderately cutinized (Fig. 4m), finely striate and consisting of ± isodiametric cells with wavy anticlines and rounded stomata ca. 35 lm long, recalling those on leaves. Pollen in situ (Figs. 4h–l). Pollen removed from stamens tricolpate, finely reticulate, 16–20 lm long and 14–18 lm in equatorial diameter (for light microscopy of pollen in situ see also Bu˚zˇek et al. 1967, pl. 3, figs. 1–4 indicating larger dimensions, i.e. 21–25 · 17– 18 lm for Markvartice; and Pacltova´ 1984, pl. 3, figs. 1–8 indicating smaller dimensions, i.e. 15–17 · 13–16 lm for the record from Borna. The size variability may be in part due to maceration process or maturity. Vegetative material. Branches bearing the simple leaves show alternate phyllotaxy with

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leaves grouped distally, (Fig. 5a–b). The petiole base encloses the axillary bud such that no traces of buds can be seen in the axil. On branches that have partly shed their leaves the buds are well preserved as seen in specimens

from Kucˇlı´ n (Fig. 5c) and Ipolytarnoć (personal observations). Twigs show closely spaced annual increments (Fig. 5b). Stipules caducous, leaving traces that surround the leaf scar.


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b Fig. 4. Staminate inflorescences of Platanus neptuni. a Globose inflorescence showing domed impressions of individual stamen connectives. Markvartice, NM G 2973. b Globose inflorescence on long stalk (note diagnostic rim at arrow). Markvartice, NM G 2960. c Mature inflorescence shedding stamens. Note conical connectives capping the elongate anthers. Markvartice, NM G 2957. d, e Globose inflorescences with in situ pollen preserved. Markvartice, NM G 2964 and Borna-Ost, MfN 9130. f Inflorescence attached to stalk with prominent circumscissile rim (arrow) and many flowers delimited by overarching perianth. Plesna´, V 146 /44–45 m, NM G 8103. g Portion of globose inflorescence three-dimensionally preserved and apparently corresponding morphologically to the impression specimens like in Fig. 4a. Plesna´,V 146 /63.5–64 m, NM G 8105. h In situ tricolpate pollen from specimen in Fig. 4d. i In situ pollen from inflorescence in g. j Pollen from inflorescence in Fig. 4f. k, l Detail from inflorescence specimen of Fig. 4d. m Epidermal structure of the connective from an isolated stamen showing striations and stomata. Markvartice, PRC MR 508/2. Scale bar in a–e ¼ 5 mm, f–g ¼ 1 mm, h–j ¼ 10 lm, k ¼ 4 lm, l ¼ 1 lm, m ¼ 50 lm

Foliage morphology (Figs. 5–7). Leaves dominantly simple in some populations, dominantely compound in other populations, the compound leaves trifoliolate, or rarely quinquefoliolate and pedately arranged; leaflets shortly petiolulate, lateral usually ± asymmetrical at base, sub-sessile, exceptionally shortly fused at the base, median symmetrical. Simple leaf and leaflet blades lanceolate to elliptic to oblanceolate, varying greatly in dimensions from 3 · 10 mm to 50 · 200 mm and more. Petiole short to medium length, expanded at the base, lamina apex acuminate or narrow to widely acute, base narrow (rarely more widely) cuneate Margin bluntly toothed except for lower part, rarely the complete lamina entire-margined (e.g. at Flo¨rsheim – Engelhardt 1911, pl. 38, Fig. 3, as Quercus elaena); teeth simple, glandular, variable in size depending on leaf forms, basal side convex, apical side straight to convex, sinuses shallowly rounded, asymmetrical. Venation semicraspedodromous to camptodromous, midrib straight, medium thick, secondaries ± regularly spaced, sub-parallel, straight to slightly curved towards the margin, typically arising from midvein at angles of about 60, in narrow forms and in more basal positions steeper, joined well within the lamina in ± angular loops, interspaced with one or two intersecondary veins, tertiaries partly parallel with secondaries or of irregular course, forming reticulate isodiametric meshes with higher-order veins, ultimate areoles often with free ending veinlets (Fig. 6b, c). Texture

coriaceous. Stipules (Fig. 6d, e) detached, fragmentary, strap-like, longitudinally striate, truncate at the base (attachment scar), frayed in shreds along veins at the top, ca. 3–4 mm wide and mostly 25 mm long (preserved as compressed, tube-like basal parts without lamina). Leaf epidermal anatomy (Fig. 8b–h). Adaxial cuticle moderately thick, smooth, rarely faintly striated along veins (SM. B 527c), in very small specimens (sun leaves?; e.g. UUG Uc 9–106) also over intercostal areas, ordinary cells polygonal-lobate, ca. 30–50 lm across, anticlines coarsely to shallowly undulate, compound trichome bases lacking or widely dispersed in small laminae (Fig. 8c). Abaxial cuticle (Fig. 8d–h) mostly smooth (finely striate in small laminae), ordinary cells polygonallobate, ca. 35–50 lm across, anticlines shallowly to markedly and deeply undulate; stomata anomocytic (to laterocytic), widely elliptic, typically 35–50 lm long, thickened peripherally, often with concentric cutin folding, stomatal ledges thickened all their length, outer stomatal aperture large elliptic, inner aperture slit-like, in best preserved specimens showing T-pieces at the poles; trichome bases sparsely distributed in larger leaves, denser in smaller leaves, compound, 28–50 ()65) lm across, formed by (1-) 2 or more basal cells and a circular attachment scar of the distal part of the trichome, the periphery of the trichome bases thickened and surrounded by radiating cutin ridges; complete glandular trichomes rarely preserved (Fig. 8d-f), with terminal parts disc-shaped, rounded to broadly elliptic,

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65–100 lm across, generally without internal cellular structure and filled with yellowish secretory matter, exceptionally preserving small isodiametric cells within the shield (Fig. 8f).

Cuticle of stipules (Fig. 8b) very thick, longitudinally striate, cells typically quadrangular, 15–25 lm wide and 25–60 lm long, isodiametric to elongate, with shallowly


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b Fig. 5. Simple leaves and twigs of Platanus neptuni mf. reussii. a Simple leaf attached to a twig with annular growth increments. Kucˇlı´ n, epitype NM G 8112, b Twig with attached simple leaves and annular growth increments, Kucˇlı´ n, coll. Valı´ cˇek, c Detail of twig from a, showing an intact bud and the abscission marks of fallen leaves and stipules. d Simple leaf showing two small lobes in apical part, Bı´ lina Mine, UUG s.n. e Leaf compression with fine dentation, Flo¨rsheim, GIK 955. f Small simple leaf with typical dentation, holotype, orig. Ettingshausen 1866, pl. 14, Fig. 4, Kucˇlı´ n, GBW 6145. g Leaf compression with coarse dentation (note enlarged base of petiole), Seifhennersdorf, MMG s.n. Scale bars ¼ 10 mm

undulate anticlines. Trichome bases scattered over the surface, composed of groups of up to 5 darker stained, smaller cells with a circular attachment scar. Stomata scattered, anomocytic, rarely with smaller cells on the periphery (±laterocytic), narrowly elliptic, 50–60 lm long, with long aperture formed by thickened ledges. Pattern corresponding to that found on the stalks and leaf petiole. Discussion Variability in leaf morphology. Three kinds of foliage are associated with the same sort of infructescences of P. neptuni: quinquefoliolate, trifoliolate and simple. Petioles are relatively short and expanded basally. It is difficult to recognize the complete leaf morphology in fragments without petiole because the venation and form of the blades of simple leaves and individual leaflets are the same except for differences in symmetry (lateral leaflets tend to be asymmetrical whereas simple leaves and terminal leaflets tend to be symmetrical). The identical epidermal structure of all three leaf forms and co-occurrence data (Table 1) reinforce our conclusion that they belong to the same species. Quinquefoliolate leaves are extremely rare but their organization is helpful in showing similarity to extant lobed species of Platanus. The pedate arrangement of the leaflets in 5-foliolate leaves (Fig. 7a, d) follows the palinactinodromous pattern of the primary venation characterising the 5-lobed leaves of Platanus subg. Platanus. The earliest epithet connected with this form is hibernica, based on Dewalquea hibernica Johnson and Gilmore (1921) from the Late Oligocene Lough Neagh

core of Ireland (Fig. 7a). A similar but fragmentary specimen of five leaflets (Fig. 7d) was recovered by Walther (1985) from the Late Eocene flora of Klausa, Germany in association with numerous trifoliolate leaves. Emphasizing the identical epidermal features, he placed both the 3- and 5-foliolate leaves within Platanus fraxinifolia (Johnson & Gilmore) Walther. Trifoliolate leaves (Fig. 7b, c, e) are generally more common than quinquefoliolate and in the Paratethys area tend to be more common than simple leaves (e.g. Poma´z – Hably 1994, Vero¨cemaros – Hably 1982, Mesta Graben – Palamarev et al. 1999). However, they occasionally co-occur in populations with predominantly simple leaves (e.g. Bı´ lina, Bo¨hlen, Kucˇlı´ n, Flo¨rsheim – Walther 1985, Kvacˇek in press, this paper Fig. 7b, c, e). The earliest epithet connected with this leaf form is fraxinifolia based on Dewalquea fraxinifolia Johnson and Gilmore (1921), also from the Lough Neagh core, Washing Bay, Ireland (National Museum, Dublin). The leaflets are as a rule free and shortly petiolulate but rarely they are shortly fused at the base (cf. Debeya hungarica Hably, Hungary – Hably 1982). There is no twig available to show the attachment of the petiole itself but the enlarged, apparently hollow, base of the petiole (Fig. 7d) suggests that it attached directly over the bud as seen in the extant subgen. Platanus. The simple-leaved form predominates at many sites (Table 1). Recently recovered twigs with attached leaves (Fig. 5a–c) from Kucˇlı´ n prove that the enlarged petiole base covers the bud like in subgen. Platanus. The earliest epithet applied to these simple leaves seems to be reussii (Myrica reussii Ettingshausen

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Fig. 6. Leaves of Platanus neptuni mf. reussii and associated stipules. a Simple incomplete leaf showing enlarged petiole base. Kundratice, NM G 7910. b, c Detailed venation and margin. Note intersecondary veins and glandular teeth. Kundratice, NM G 4638. d Stipule (Equisetites ettingshauseni Engelhardt 1898, pl. 1, Fig. 31), Suletice-Berand, MMG SuBe 236d. e Stipule, Kundratice, ML 208.1. Scale bar a ¼ 10 mm, b–e ¼ 5 mm

1866 from Kucˇlı´ n); at least we are not aware of any other name published earlier, which might have priority. A single aberrant specimen from the Bı´ lina Mine (Fig. 5d) shows two slightly enlarged teeth on either side of the leaf apex suggesting a transitional form between trifoliolate and simple forms. The list of localities (Table 1) shows the overlapping distribution of the leaf forms described above and their co-occurrence with specified reproductive organs. At one site (Klausa), an infructescence (Fig. 2c) was found together with only compound leaves (Fig. 7d, f, Walther 1985). At Kucˇlı´ n several infructescences (including the species lectotype – Fig. 2g) and inflorescences are available, and both simple (Fig. 5a–b) and trifoliolate (Fig. 7b–c) leaves are present. At all other localities, the inflorescences are found mainly in association with simple leaves or just isolated (e.g. Fasterholt area). The Markvartice locality, which provided some of the most informative staminate and pistillate inflorescences, yielded mostly

fragmentary foliage, but the laminar asymmetry of some specimens (e.g. Fig. 7g) suggests that they partly represent compound leaves. The simple and compound leaves of this complex have been traditional treated separately as individual morpho-species. In our opinion, however, they are more properly regarded as mere leaf forms of a single species. The shared characters of the laminae in simple and compound leaves are: elliptic to oblong lanceolate outline, long cuneate base, acuminate to blunt-acute apex, semicraspedodromous venation usually with intersecondaries (contrary to P. bella – Kvacˇek et al. 2001) and the margin which typically, at least in the upper part, is ± bluntly serrate to denticulate with glandular teeth that are convex on their basal sides and straight to concave on the apical sides, with shallowly rounded sinuses. Sun and shade leaves. The leaf epidermal structure (Fig. 8c–h) does not vary much in correlation to the dimensions of laminae, but the smallest forms, presumed to be sun leaves,


15

Fig. 7. Compound foliage of Platanus neptuni mf. hibernica and mf. fraxinifolia. a Complete leaf of five leaflets, holotype of Platanus neptuni mf. hibernica, Lough Neagh core, GLAHM Pb 2351. b, c Trifoliolate leaves of Platanus neptuni mf. fraxinifolia, Kucˇlı´ n, MMG KIN 84 and 423:1:22. d Leaf base of Platanus neptuni mf. hibernica showing enlarged petiole base, orig. Mai and Walther 1985, pl. 13, Fig. 5, Klausa, MMG KL 247. e complete trifoliolate leaf of Platanus neptuni mf. fraxinifolia with well preserved venation, Bı´ lina Mine, NM G 7844a. f Trifoliolate leaf of Platanus neptuni mf. fraxinifolia, orig. Mai and Walther 1985, pl. 13, Fig. 4, Klausa, MMG KL 142. g Leaflet (?) with asymmetrical base and typical dentation, Markvartice, UUG MR 1044. Scale bars ¼ 10 mm

have less pronounced undulation of anticlines, and glandular trichome bases very densely distributed on the abaxial side and quite dense on the adaxial side with fine striations over

intercostal areas on both adaxial and abaxial leaf epidermides. At the other extreme, abnormally large leaves presumed to be shade leaves, typically have large, X-shaped undulations on

16



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b Fig. 8. Epidermal structure of Platanus neptuni. a Peduncle of pistillate infructescence (horizontally oriented). Markvartice, PRC MR 508/1. b Basal part of stipule (horizontally oriented). Markvartice, PRC MR 59. c Adaxial leaf side of smaller type with trichome bases. Plesna´, PRC V 146-3/2. d Abaxial leaf side on midrib (horizontally oriented) showing stoma and glandular trichome. Plesna´, PRC V 146–268. e, f Abaxial side of submacerated leaf fragment with stoma and glandular trichome in interference and phase contrast light. Plesna´, PRC V 146–268. g, h Adaxial and abaxial sides of ordinary simple leaf type. Flo¨rsheim, SM. B 314b/1. Scale bars ¼ 50 lm

both leaf surfaces, and fewer trichome bases. The striation in such leaves is almost absent, faintly seen over the abaxial side of the midrib (Kvacˇek and Walther 1978). The form of stomata and compound trichome bases are stable diagnostic characters, and are recognizable even on very poorly preserved cuticles (see Kvacˇek in press). Stipules. The structures that Bu˚zˇek et al. (1967) interpreted as detached stipule remains are similar to stipule bases observed on the cultivated young trees of P. kerrii (Komarov Botanical Institute, St. Petersburg). In P. kerrii, the basal part of the stipule, which forms a tube around the stem, persists longer than the laminar part of the stipule, and falls off only after the laminar part of the stipule decays. Comparison of reproductive organs. The well-preserved specimens described and illustrated as P. neptuni from the Miocene Fasterholt flora of Denmark by Friis (1985, see also Koch et al. 1973, pl. 2, Fig. 21) contribute to our concept of the species. The Fasterholt infructescence axes are elongate ellipsoidal or ovoidal, and bear 4 to 8 achenes per flower. The stalk of the infructescences is (according to Friis 1985, p. 29) 1.9–4.0 mm wide and up to 15 mm long, with what appears to be a distinct joint between the stalk and fruiting head. Associated achenes are unilocular, narrowly triangular or obovate, tapering into a persistent and slightly curved style. A longitudinal slit runs from the top of the style to about 1.5 mm below the style base. The length of the achene without style is 6.0–8.5 mm, max. width 0.6–0.9 mm. Friis (1985) observed that the locule occupies the lower two thirds of the achene, which is also seen in the Kundratice specimens (Fig. 3g), and is lined by a layer of transversely elongate sclereids. The outer

wall of the achene is composed of narrow, longitudinally aligned sclereids with finely pitted walls. The outer epidermal cells are thin-walled and mainly tabular with straight to anticlinal walls; and dotted with simple, equiaxial or longitudinally elongate hair bases 0.02 to 0.1 mm long composed of 1–4 slightly enlarged cells with thickened walls and arranged in distinct longitudinal rows. The hair bases are closely spaced, about 5 per 0.1 mm2. Seeds are 3.5–4.5 mm long, 0.8–1 mm wide, hemiorthotropous and pendant. Staminate receptacular cores are elongate-ellipsoidal 1.5–6.5 mm long, and 0.9–3.3 mm wide, borne on a thick stalk, 1.4–3.0 mm broad (Friis 1985, pl. 5, Fig. 1). Friis indicated that there are ‘‘no attached anthers, but remnants of flowers show 5–8 central small protrusions, probably representing reduced carpels, surrounded by 6–7 elongate scars that probably represent the stamens’’. In comparison with the lectotype and associated specimens of P. neptuni from Kucˇlı´ n, the Fasterholt material shows both similarities and differences. Similarities include the relatively thick stalk, both on the staminate inflorescences and the infructescences, with a thickened rim below the head, and the slightly transversely elongate organization of epidermal cells on the stalks (in contrast to the longitudinally elongate epidermal cells of peduncles in the extant species). The isodiametric to slightly transversely elongate epidermal cells of the stalk are also observed in the Markvartice specimens (Fig. 8a). Friis observed that the ‘‘infructescence axes’’ (receptacular cores) are elongate ellipsoidal or ovoidal, and staminate ‘‘axes’’ elongate ellipsoidal. This elongate, rather than globose, shape is sometimes observed in the Czech material, as well.

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The Fasterholt fossils apparently differ from the Czech specimens by the number of carpels per flower: 4–8 in Fasterholt specimens vs. 3 in Kucˇlı´ n and Plesna´. The isolated achene illustrated by Friis is narrowly triangular or obovate, like some of the specimens we studied (e.g. Fig. 3f–h). Judging according to our material, the pistillate immature inflorescences might not disintegrate easily because detached achenes and cores devoid of achenes were not recovered at most localities of unsieved impression/compression material. Not all records of head-like inflorescences belong to Platanus neptuni. Those recently described under this name by Gu¨mbel and Mai (2002) from the Miocene of Rho¨n differ in straight fine filiform styles and do not bear any sign of glandular trichomes (exocarp cuticle preparations by Z.K. at MfN). We consider them inflorescences of Sparganium similar to Sparganium valdense Heer. Nomenclature. At the time of the first revision (Bu˚zˇek et al. 1967) it was decided to employ the epithet neptuni for all associated organs including staminate and pistillate inflorescences, infructescences, simple leaves and stipules because of the shared epidermal anatomy. Walther (1985) treated the populations with compound leaves from Washing Bay and from Klausa as a separate species P. fraxinifolia, yet Mai (in Mai and Walther 1985) treated co-occurring infructescences from the same basin and levels as P. neptuni. We know now that the simple and compound leaves often occur together in different proportions associated with a single type of infructescence and there is little doubt that they belong to the same plant because of the repeated co-occurrence and shared epidermal anatomy. The nomenclature of the P. neptuni complex can be solved in different ways. We prefer to maintain the name P. neptuni in a broad sense for the whole plant and distinguish infraspecific morphotaxa as forms including reproductive organs (morphoforma neptuni – autonym) and various kinds of foliage (mf. hibernica for quinquefoliolate specimens, mf. fraxinifolia for trifoliolate specimens and mf. reussii for simple

leaves). Another approach would be to retain distinctions at the species level, but the morphological intergradation and uniform epidermal anatomy support their treatment as a single species. The system of classification of the Platanaceae will certainly require revision taking into consideration more of the extinct entities as well as the extant species. Numerous other Cretaceous and Tertiary platanaceous taxa are known in only a limited way from isolated leaves, wood, or reproductive structures. Improved understanding of their whole-plant morphology will likely influence our understanding of the position and rank of Platanus neptuni within the complex. At the present state of our knowledge, we prefer, mostly for pragmatic reasons, to maintain the assignment of our P. neptuni complex to the extinct subgenus Glandulosa of Platanus, not treating it collectively as a distinct genus nor splitting it into several morpho-genera for individual detached organs. The available names, e.g. Celastrophyllum (for simple leaves), Debeya (for trifoliolate) and Dewalquea (for quinquefoliolate) designate purely artificial entities, requiring reconsideration for phylogenetic analyses (see also Kvacˇek et al. 2001). As specified below, one of the original syntypes of Sparganium neptuni (Ettingshausen 1866, pl. 7, Fig. 11F, our fig 2G) was selected as the lectotype of P. neptuni (Kvacˇek et al. 2001) and another syntype was illustrated as the paratype (Hably et al. 2001 , pl. 34, Fig. 3). It is worth noting that one of the originally figured syntypes (Ettingshausen 1866, pl. 7, Fig. 9) is actually from another locality, Priesen, in Czech Brˇ ezˇańky (mislabelled in figure caption) and represents an infructescence of Liquidambar (Hably et al. 2001, p. 37). The leaf remains originally included in the same species of Sparganium (Ettingshausen 1866, pl. 7, figs. 16, 18) have been excluded as unrelated monocot foliage (Bu˚zˇek et al. 1967). The remaining syntypes are accepted as additional examples of Platanus neptuni, although the specimen in Ettingshausen’s pl. 7, Fig. 14 is missing.


Conspectus of the Platanus neptuni complex. Family Platanaceae Dum., genus Platanus L. subgen. Glandulosa Z. Kvacˇek, Manchester & Guo, Int. J. Plant. Sci. 162: 444. 2001. Platanus neptuni (Ettinsgshausen) Bu˚zˇek, Holy´ & Z. Kvacˇek, Monatsber. Dtsch. Akad. Wiss. 9: 205. pl. 1. figs. 1–4, 6 (non 5 ¼ Sloanea artocarpites), pls. 2–4. 1967 (basionym: Sparganium neptuni Ettingshausen, separatum, Denkschr. K. Akad. Wiss. math.-nat. Cl. 26: 31. pl. 7. figs. 10–16 (non figs. 9, 17–18). 1866 [lectotype selected by Kvacˇek et al. 2001: 453, Fig. 7 A – No. BP 55.2491.1 (Ettingshausen 1866, pl. 7, Fig. 11) re-figured in Hably et al. 2001, pl. 33, Fig. 5 and in this paper Fig. 2g; paratype selected and re-figured by Hably et al. 2001: 37, pl. 34, Fig. 3 – No. BP 55.2337.1 (Ettingshausen 1866, pl. 7, Fig. 13)]. morphoforma neptuni (Bu˚zˇek et al. p. 205. pl. 2. figs. 6a–b, 7 a–b, 8 a–b, 9. pl. 3. figs. 5–9. 1967. Type locality Kucˇlı´ n, N Bohemia, Late Eocene. Globular infructescences/staminate & pistillate inflorescences, dispersed achenes, stamens). morphoforma reussii (Ettingshausen) stat. & comb. n. (basionym Myrica reussii Ettingshausen, separatum, Denkschr. K. Akad. Wiss. math.-nat. Cl. 26: 44. pl. 14. Fig. 4. 1866 – holotype No. 6145, Geologische Bundesanstalt, Wien – re-figured in Fig. 5 f, epitype selected here – No. NM G 8112, figured in Fig. 5a. Type locality Kucˇlı´ n, N Bohemia, Late Eocene. Simple leaves). morphoforma fraxinifolia (Johnson & Gilmore) stat. & comb. n. (basionym Dewalquea fraxinifolia Johnson & Gilmore, Sci. Proc. Roy. Soc. Dublin, 16: 327. pl. 11. figs. 4–11. pl. 12. Fig. 3. text-Fig. 3. 1921.Type locality Lough Neagh core, Ireland, Late Oligocene. Trifoliolate leaves). morphoforma hibernica (Johnson & Gilmore) stat. & comb. n. (basionym Dewalquea hibernica Johnson & Gilmore, Sci. Proc. Roy. Soc. Dublin, 16: 326. pl. 11. figs. 1–3. pl. 12. figs. 1–2. 1921 – lectotype selected here No. Pb 2351, Hunterian Museum, Glasgow, figured in Johnson and Gilmore 1921, pl. 11,

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Fig. 1 – re-figured in Fig. 7a. Type locality Lough Neagh core, Late Oligocene. Quinquefoliolate leaves). Comparison with other Platanaceae. Known from inflorescences, fruits, pollen, twigs, and leaves, ‘‘the Neptunus Plane’’ is one of the most completely reconstructed extinct dicotyledons. The affinity with Platanus was not recognized by pioneering paleobotanists and continues sometimes to be challenged to present day (e.g. Palamarev and Staneva 1995). Therefore, it may be useful to reiterate the morphological and anatomical justification for a confident position within the Platanaceae. Both reproductive and vegetative structures independently conform to extant members of the Platanaceae. Reproductive features diagnostic of the family include globose staminate and pistillate inflorescences composed of numerous few-fruited flowers, folicular fruits with tapering persistent styles, stamens with short filaments, elongate anthers and prominent connectives, tricolpate microreticulate pollen. The leaves are consistent in alternate phyllotaxy and the anomocytic (to cryptolaterocytic) stomatal type (Kvacˇek et al. 2001) with all extant Platanaceae. The lamina shape (unlobed and partly compound) and margin (serrate without scalloped sinus) are unlike most extant species yet similar to the subtropical/tropical Asian species, Platanus kerrii. At first it was thought that P. neptuni might be most closely related to the extant P. kerrii of subgen. Castanophyllum Leroy because they both have unlobed finely serrate elliptical leaves (and slightly smaller pollen than the mean of the genus – cf. Crane et al. 1988). However, the fossils are distinguished by prominent, peltate glandular trichomes, deeply undulate anticlines of epidermis (Kvacˇek 1970) and solitary rather than multiple heads per peduncle. We stress that P. neptuni shares other characters with subgenus Platanus, such as inflorescences apparently solitary, palinactinodromous organization of 5-foliolate leaves (like the primary venation of 5-lobed leaves) and the enlarged petiole base covering the bud

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(Fig. 5a–c). Because of their enclosed position within the petiole base, the buds cannot develop without shedding the leaves. From this we infer that the foliage of P. neptuni was deciduous, as is also suggested by the abundance of the foliage at many fossil sites. In the single evergreen species, Platanus kerrii, the axillary buds are not enveloped by the petioles (Leroy 1982). Some features of P. neptuni are not seen in any extant Platanus: tendency for compound leaves, the swollen rim at the top of staminate and pistillate inflorescence stalks, and abundant conspicuous glands on foliage and fruits. Kvacˇek et al. (2001) stressed these features to support recognition of the subgenus Glandulosa, which is typified by the additional Paleocene species P. bella that is known from tri- and rarely quinquefoliolate leaves. Platanus neptuni is distinguished from it by subtle details of venation (regular intersecondary veins) and cuticular structure (coarser undulation, scarcity of striation). Friis (1985) observed that the internal sclerenchyma cells of the achenes of P. neptuni are transversely aligned, rather than longitudinally aligned in modern species of Platanus – this might be another feature diagnostic of subgenus Glandulosa. No other Platanaceae, fossil or modern, have a swollen circumscissile rim on the stalk below the inflorescences. Studying specimens from the Fasterholt flora, Friis (1985) was the first to call attention to this feature, which she referred to as a joint between the head and peduncle. It was apparently not abscising at this position, however, because we have not encountered any specimens showing disarticulation from the stalk. Such a rim is absent, for example, from Macginicarpa Manchester (1986) and other Early Tertiary and Cretaceous records (e.g. Pigg and Stockey 1991, Crane et al. 1993, Maslova 2002). The rim is not divided into a whorl of individual scars, which would be expected if it represented the position of fallen bracts. Rather, it forms a continuous band around the base of the head. Cuticle preparations do not suggest that it was nectariferous because there are no secretory struc-

tures nor concentration of adhering pollen and other particles. Possibly, this rim may represent a scar left by the deciduous petiole that enveloped the reproductive structures in bud stage. This interpretation would suggest that both vegetative and reproductive buds were covered by the petiole of the previous season’s leaf, as in extant subgenus Platanus, but that the buds were stipitate, or stalked, unlike those of any living Platanus species. We conclude that the Platanus subgen. Glandulosa members represent an extinct lineage of the family without modern descendants. The elliptical unlobed leaves probably evolved convergently with Platanus kerrii. Platanus neptuni may be secondarily simple (with compound-leaved ancestry), whereas P. kerrii may derive from a lineage that had simple leaves from its inception. In addition, the kind of compound leaf produced by members of subgen. Glandulosa, in which each of the leaflets normally is unlobed, is easily distinguished from those of the extinct genera Platanites Forbes and Erlingdorfia K. Johnson (1996), in which the terminal and sometimes the lateral leaflets are markedly lobed. Palaeoecology and sociology. The earliest reliable records of Platanus neptuni come from the Upper Eocene of Central Europe. Most recently described local floras from the Zeitz Floral Assemblage (‘‘Florenkomplex’’) of Saxony (Mai and Walther 1985) indicate very equable subtropical climate with high annual precipitation. The plant assemblage of Klausa, where the trifoliolate leaf form predominates, occurs in a clay layer within fluviatile sands of the palaeodrainage known as the ‘‘ZwickauAltenburg River’’. The flora includes, besides Platanus neptuni, many typical Late Eocene elements, such as Rhodomyrtophyllum, Eotrigonobalanus, Daphnogene, Steinhauera, Doliostrobus, Sloanea, Retinomastixia, Gordonia, etc. (Mai and Walther 2002, pp. 15–17). The community can be called a humid subtropical laurel-conifer forest. The Late Eocene Stare´ Sedlo Formation from the adjacent Bohemian part of Czech Republic, which is an extension of the Zeitz complex, contains a very similar


Notophyllous Broad-leaved Evergreen Forest dominated similarly by evergreen Lauraceae, Fagaceae, Myrtaceae, Theaceae and Hamamelidaceae. The sedimentary setting of the fossiliferous deposit suggests riparian (azonal) forest vegetation and the representation of P. neptuni is negligible relative to the quantity of collected specimens of other arboreal components. The Late Eocene site of Kucˇlı´ n in North Bohemia, studied initially by Ettingshausen, is not yet well revised. According to the recent research, the vegetation is mostly evergreen (Lauraceae, Sloanea) or semi-evergreen (Cedrelospermum, Ziziphus). The physiognomy of the assemblage may suggest sub-humid subtropical climate. This volcanic lake was probably surrounded by mesophytic forest vegetation on slopes with thermophilic conifers (Doliostrobus, Tetraclinis) and broadleaved elements (Engelhardia, Sloanea, and Hooleya) and palms (Sabal). The recently suggested reconstruction of the Late Eocene landscape development in North and West Bohemia interprets the volcanic (Kucˇlı´ n, core Hlinna´ and others) and fluviatile facies (Stare´ Sedlo Fm.) as contemporaneous (Kvacˇek 2002). Near-shore environmental conditions were later connected with the Early Oligocene Tard Clay assemblages in the Hungarian part of the Paratethys Province. Besides the common P. neptuni complex many thermophilic plants survived from Late Eocene, as Doliostrobus, Tetraclinis, Laurophyllum sp. div., Sloanea, Engelhardia, and Eotrigonobalanus. Only some components of this flora are new immigrants – e.g. Ailanthus, Calocedrus, Craigia, Quercus lonchitis, which penetrated at that time even further north. In some other Oligocene sites of southern Europe, namely in southern Bulgaria and Hungary, the trifoliolate form predominates (Palamarev et al. 1999; Hably 1982, 1994). In the Tethys-Paratethys province deciduous broad-leaved elements are during Early Oligocene very scanty (Platanus schimperi, Alnus) while Leguminosae are common and well diversified. The Boreal Province of central Europe is dominated in the very early

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Oligocene by deciduous broad-leaved elements (Haselbach and Kundratice Floral Assemblages – Kvacˇek and Walther 2001), where only a few thermophilic elements, such as Tetraclinis, Lauraceae, and Sloanea, persisted within the predominantly deciduous forests. The relative abundance of the P. neptuni complex differs greatly there, depending on the environment. In the basinal assemblages, dominated by Eotrigonobalanus and other riparian – swamp elements, remains of P. neptuni are very scanty. On the other hand, at more or less mesophytic ‘‘volcanic’’ sites (e.g. Kundratice, Suletice) they are much richer. The German locality Seifhennersdorf at the Bohemia – Saxony border is in this respect aberrant, having a good representation of azonal trees, e.g. Taxodium, Alnus, as well as P. neptuni (Walther 1996). Markvartice in north Bohemia (Bu˚zˇek et al. 1976), probably of late Early Oligocene age, seems to be the richest occurrence of all kinds of organs belonging to P. neptuni, accompanied with the Mixed Mesophytic Forest vegetation, including Sloanea, Liriodendron, Lauraceae, Betulaceae and Oleaceae. The newly revised Early Oligocene flora at Flo¨rsheim in the Mainz Basin (Kvacˇek in press) is another rich occurrence of P. neptuni connected with humid subtropical Nothophyllous Evergreen forest. The physiognomical similarity of the Flo¨rsheim and the Tard Clay assemblages is indicated by shared elements, like Ceratozamia, Sloanea, Eotrigonobalanus, but there are many more new immigrants at Flo¨rsheim typical of mastixioid Miocene floras, e.g. Laurus abchasica, Quercus bavarica, Gordonia hradekensis etc. Subtropical to warm temperate climatic conditions can be expected for the whole Boreal Province during the Oligocene with some fluctuation of mean annual temperature and differences of precipitation regime due to palaeogeographical position. Abundant and rich Late Oligocene (Egerian) occurrences of P. neptuni are connected with the Paratethys area. Here again, evergreen broad-leaved forest vegetation predominated, with admixture of some more

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deciduous elements, e.g. Alnus, Ulmus and Acer (Andrea´szky 1966, Kvacˇek and Hably 1991). At some sites, the fraxinifolia leaf form is well represented, e.g. Janda – Fruska Gora (Mihajlovic 1996–1997), and Vero¨cemaros (Hably 1982), always connected with more thermophilic vegetation than in other Egerian sites. The other Late Oligocene occurrences of P. neptuni in Central Europe (e.g. Thierbach Floral Assemblage, Kleinsaubernitz core – Mai and Walther 1991, Walther 1999) are much more scanty, connected with more or less warm temperate Mixed Mesophytic type of vegetation. Early Miocene sites are similarly differentiated geographically. The most abundant occurrence of the P. neptuni complex was described from Ipolytarnoć (Hably 1985a) and adjacent Lipovany (Neˇmejc and Knobloch 1973, Sita´r and Kvacˇek 1993). Assemblages of both the sites are dominated by thermophilic vegetation including, besides P. neptuni, also Lauraceae, Leguminosae, Engelhardia, Pungiphyllum cruciatum, and palms. The corresponding floras in the Bohemian Massif (Bı´ lina, Cypris Shale) contain much higher proportion of deciduous elements, indicating that they correspond to the warm temperate Mixed Mesophytic forests. Platanus neptuni is not frequent there. During the Middle and Late Miocene, P. neptuni became a relic, withdrawing southeastwards to Romania and Bulgaria (Kvacˇek 1994, Kovar-Eder et al. 1996). Judging from the dominant elements in the richest occurrences of the P. neptuni complex, there is no doubt that this plant was thermophilic, probably with nearly the same climatic requirements as extant Platanus species growing in southern Mexico or P. kerrii from Viet Nam and Laos, although it apparently tolerated colder situations as well. As stated in the descriptive part, P. neptuni shed foliage like the Mexican plane tree species and differs from P. kerrii in which the bud is not completely covered by the leaf petiole. In its native area of Laos and north Viet Nam, P. kerrii is evergreen, bound to still warmer more or less tropical conditions. As Walther (1985) stressed, P. neptuni had broad ecological

amplitude having been more often connected with zonal (volcanic and near-shore marginal) rather than alluvial – basinal habitats. Until recently, P. neptuni was considered a European endemic (Hably et al. 2000). Our inspection of the collections from western Oregon, USA, housed in the Smithsonian Institution, revealed a new occurrence of this element in the Late Oligocene Yaquina flora (McClammer 1978, Manchester, Hickey, and Kvacˇek in progress). At this site, only the trifoliolate leaf form is represented, associated with both pistillate and staminate inflorescences. These inflorescences have the typical circumsissle scar on the peduncle beneath the head. The plant assemblage of Yaquina is connected with shallow marine sandstone/siltstone facies. It is dominated by conifers, such as Pinus, Pseudolarix (Gooch 1992), taxodioids, Chamaecyparis and broad-leaved Betulaceae with additional Acer, Hydrangea, Cornus, Smilax and Leguminosae among other unidentified dicots. It is likely that the P. neptuni populations with the prevailing trifoliolate foliage may indicate Eocene or Oligocene exchange across the Atlantic or these may be independent parallel occurrences that are remnants of the widespread high latitude Paleocene species, P. bella. This case is another example of the disjunct distribution of Tertiary plants between North America and Europe, as in Tetraclinis salicornioides (Kvacˇek et al. 2000). Appendix Critical records of the Platanus neptuni complex in Europe including previous misidentifications and synonyms (numbers in [] refer to localities in table 1; references on earlier publications with comma (,) after the taxon name in case of its first publication, with dash (–) in other cases). Unger 1850 [18]: Panax longissimum, p. 44, partim, pl. 24, fig. 23 (non fig. 22 ¼ lectotype designated here). Ettingshausen 1866 [9a]: Sparganium neptuni, p. 31, partim, pl. 7, figs. 10–15; Myrica reussii, p. 44, pl. 14, fig. 4.

Z. Kvacˇek and S. R. Manchester: Platanus neptuni complex from the Tertiary of Europe Saporta 1867 [2]: Quercus singularis, p. 68, pl. 5, fig. 5; Fraxinus juglandina, p. 89, pl. 7, fig. 8, pl. 9, figs. 13–15. Ettingshausen 1869 [9a, g]: Ceratopetalum bilinicum, p. 6, pl. 40, figs. 26, 30–31; Ceratopetalum haeringianum – p. 6, pl. 40, figs. 27–28, pl. 41, figs. 4–5; Bombax charisiaefolium, p. 11, pl. 42, figs. 4–5; Elaeocarpus europaeus, p. 16, partim, pl. 43. fig. 7; Ternstroemia bilinica, p. 17, partim, pl. 47, figs. 9(?), 10; Euonymus radobojana – p. 29, pl. 48, fig. 8; Maytenus europaea, p. 31, pl. 48, figs. 10–12; Celastrus deucalionis, p. 33, pl. 48, fig. 15; Elaeodendron degener, p. 37, partim, pl. 49, fig. 8; Hippocratea bilinica, p. 38, pl. 49, figs. 12–14; Rhus hydrophila – p. 49, pl. 51, fig. 3; Amygdalus bilinica, p. 55, partim, pl. 53, fig. 23; Cunonia bilinica, p. 64, pl. 55, fig. 21. Pilar 1883 [19]: Bombax neptuni – p. 92, pl. 13, fig. 13. Engelhardt 1885 [9d]: Amygdalus pereger – p. 71, pl. 19, fig. 3; Ardisia myricoides – p. 42, pl. 9, fig. 23; Bombax chorisiaefolium – p. 50, pl. 11, fig. 7; Carya elaenoides – p. 67, partim; Ceratopetalum bilinicum – p. 48, pl. 10, fig. 27; Ceratopetalum cundraticiense, p. 48, pl. 11, fig. 2; Ceratopetalum haeringianum – p. 49, pl. 10, fig. 24; Cunonia bilinica – p. 47, pl. 10, fig. 29; Juglans bilinica – p. 65, partim, pl. 16, figs. 23, 29, pl. 17, fig. 6; Myrica acuminata – p. 20, pl. 2, fig. 9; Myrica hakeaefolia – p. 20, partim, pl. 2, fig. 2, figs. 4, 6, 27; Myrsine clethrifolia – p. 40, pl. 9, figs. 10, 11; Quercus mediterranea – p. 22, pl. 2, fig. 33; Sparganium valdense – p. 17, pl. 2, fig. 2. Engelhardt 1898 [9e]: Belangera obtusifolia – p. 28; Benzoin paucinerve – p. 19; Bombax chorisiaefolium – p. 28; Celastrus acherontis – p. 32; Celastrus deucalionis – p. 32; Celastrus hippolyti – p. 32; Ceratopetalum bilinicum – p. 28; Ceratopetalum cundraticiense – p. 27; Ceratopetalum haeringianum – p. 27; Cunonia bilinica – p. 28; Eugenia haeringiana – p. 38; Hippocratea bilinica – p. 34; Juglans bilinica – p. 9, 35, pl. 1, fig. 24; Maytenus europaea – p. 33; Myrica acuminata – p. 13; Myrsine clethrifolia – p.24; Persoonia daphnes – p. 21; Prunus bilinica – p. 9, pl. 1, fig. 24; Quercus lonchitis – p. 15; Equisetites ettingshausenii, p. 85, pl. 9, figs. 6, 8, 11, 30, 31, 36, 37. Engelhardt 1911 [3]: Laminarites latus, partim, p. 316, pl. 37, fig. 7; Myrica lignitum – p. 327, pl. 37, fig. 43; Myrica sagoriana – p. 328, pl. 37, fig. 34; Quercus neriifolia – partim, p. 330, pl. 38, fig. 5; Quercus elaena – p. 330, pl. 38, fig. 3; Quercus argute-serrata – p. 331, pl. 38, fig. 17;

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Quercus chlorophylla – p. 331, pl. 38, fig. 12; Quercus decurrens – p. 332, pl. 38, fig. 21; Quercus lonchitis – p. 332, pl. 38, fig. 19; Quercus gmelinii – p. 333, pl. 38, fig. 35; Ulmus longifolia – p. 335, pl. 38, fig. 10; Ficus multinervis – p. 336, pl. 38, figs. 23, 27; Ficus jynx – p. 336, pl. 38, fig. 24; Ficus tenuinervis – p. 338, pl. 38, fig. 39; Salix varians – p. 342, pl. 38, fig. 45, pl. 39, fig. 1; Salix denticulata – p. 343, pl. 39, fig. 2; Laurus primigenia – p. 345, pl. 39, fig. 22; Elaeagnus acuminata – p. 356, pl. 40, fig. 27; Banksia haeringiana – partim, p. 358, pl. 40, fig. 27;Apocynophyllum amsonia – p. 362, pl. 40, fig. 41; Myrsine centaurorum – p. 366, pl. 41, fig. 11; Myrsine clethrifolia – p. 367, pl. 41, fig. 12; Myrsine zaddachii – p. 367, pl. 41, fig. 22; Sapotacites minor – p. 369, pl. 41, fig. 26; Sapotacites daphnes – p. 369, pl. 41, fig. 45; Sapotacites tenuinervis – p. 369, pl. 41, fig. 30; Bumelia ambigua – p. 369, pl. 41, fig. 28; Bumelia oreadum – p. 370, pl. 41, 27; Sideroxylon putterlickii – p. 370, pl. 41, fig. 34; Diospyros brachysepala – p. 371, pl. 41, fig. 7; Vaccinium acheronticum – partim, p. 372, figs. 18, 23; Andromeda protogaea – partim, p. 373, pl. 41, fig. 35; Panax longissimum – p. 374, pl. 41, fig. 43; Ceratopetalum haeringianum – p. 375, pl. 41, fig. 38; Ceratopetalum radobojanum – p. 375, pl. 41, fig. 48; Weinmannia sotzkiana – p. 375, pl. 41, fig. 51; Bombax chorisiaefolia – p. 377, pl. 41, fig. 57; Bombax neptuni – p. 377, pl. 41, fig. 61; Sapindus densifolia – p. 379, pl. 41, fig. 63; Celastrus persei – p. 379, pl. 42, fig. 1; Celastrus andromedae – p. 379, pl. 42, fig. 2; Celastrus acherontis – p. 380, pl. 42, fig. 4; Celastrus elaenus – p. 380, pl. 42, fig. 4; Celastrus europaeus – p. 380, pl. 42, fig. 6; Celastrophyllum myricoides – p. 381, pl. 42, fig. 12; Elaeodendron sagorianum – p. 382, pl. 42, fig. 31; Elaeodendron degener – p. 382, pl. 42, fig. 31; Euonymus heerii – p. 383, pl. 42, fig. 13; Euonymus latoniae – p. 383, pl. 42, fig. 38; Ilex dianae – p. 383, pl. 42, fig. 9; Rhamnus gaudinii – p. 385, pl. 42, fig. 9; Rhamnus rossmaessmeri – p. 386, pl. 42, fig. 33; Rhamnus eridani –p. 387, pl. 42, fig. 37; Juglans acuminate – p. 387, pl. 42, fig. 35; Juglans vetusta – p. 388, pl. 42, fig. 51; Juglans bilinica – p. 388, pl. 42, figs 1, 2; Carya heerii – p. 389, pl. 43, fig. 5; Pterocarya denticulate – p. 389, pl. 43, fig. 4; Rhus sagoriana – p. 390, pl. 42, fig. 27; Myrtophyllum caryophylloides, p. 393, pl. 43, fig. 7; Amygdalus pereger – p. 394, pl. 43, fig. 13; Amygdalus persicifolia – p. 395, pl. 43, fig. 20; Ceratonia septimontana – p. 397, pl. 43, fig. 40; Cassia berenices – partim, p. 398, pl. 43, fig. 19; Cassia

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phaseolites – partim, p. 399, pl. 43, fig. 40; Cassia cordifolia – p. 400, pl. 43, fig. 39; Alnus kefersteinii – p. 402; Fraxinus denticulata – p. 402. Johnson and Gilmore 1921 [1]: Dewalquea hibernica, p. 326, pl. 11, figs. 1–3, pl. 12, figs. 1, 2, text-fig. 1; Dewalquea fraxinifolia, p. 327, pl. 11, figs. 4–11, pl. 12, fig. 3, text-fig. 2; Dewalquea denticulate, p. 327, pl. 11, figs. 12–14, pl. 12, fig. 4, text-fig. 3. Weyland 1948 [4]: Liquidambar europaea – p. 125, pl. 19, fig. 4, Pruniphyllum rhoefolium – p. 129, pl. 20, fig. 10. Andreańszky and Nova´k 1957 [23b]: Cunonia oligocaenica, p. 47, pl. 2, fig.5; Elaeodendron oblanceolatum, p. 52, pl. 3, fig. 12. Knobloch 1961 [10c]: Alnus sp. – p. 298, partim, pl. 2, fig. 3; Comptonia sp. – p. 258, pl. 3, fig. 9 left; Ceratopetalum bilinicum – p. 277, pl. 10, fig. 12. Mai 1963 [10b]: ‘‘Sparganium neptunii – Ett.’’, p. 82, pl. 12, figs. 4–5. Andreańszky 1965 [23a]: Cunonia oligocaenica – p. 13, text-fig. 3. Andreańszky 1966 [23c]: Cunonia oligocaenica – p. 86, text-fig. 77. Bu˚zˇek, Holy´ and Kvacˇek 1967 [8d, 9a, d, e, 10a]: Platanus neptuni, pp. 203–215, pl. 1, figs. 1–4, 6 (non 5), pls. 2–4. Knobloch 1969 [14]: cf. Fraxinus sp. – p. 18, pl. 5, fig. 3, Cupania neptuni – p. 18, pl. 3, fig. 11. Kvacˇek 1970 [9e]: Platanus neptuni, p. 435, pl. 56, fig. 5 (non 1). Koch et al. 1973 [5]: Platanus sp. – p. 41, pl. 1, fig. 12; Aracistrobus sp. – p. 41, pl. 2, fig. 21. Neˇmejc and Knobloch 1973 [22a]: Engelhardtia sp. – p. 702, pl. 4, fig. 5, Laurophyllum cf. heerii – p. 711, pl. 2, figs 4–6. Knobloch 1973 [2]: Platanus neptuni – p. 283, text-figs. 1– 2. Walther 1974 [10b]: Platanus neptuni – p. 149, pls. 7– 8. Knobloch 1975 [14]: Platanus neptuni – p. 547. Givulescu 1975a [24a]: Engelhardia mirabilis, p. 163, pl. 2, figs. 1–6, pl. 8, figs. 15–17. Givulescu 1975b [24a]: Engelhardia mirabilis, p. 73, text-fig. 2/6. Givulescu 1976 [24b]: Engelhardia mirabilis – text-figs. 3–5. Bu˚zˇek, Holy´ and Kvacˇek 1976 [10a]: Platanus neptuni – p. 94. Pantiæ and Mihajlovic [26] 1977: Amygdalus radobojana – p. 161, pl. 4, fig. 2.

Mai and Walther 1978 [7c]: Platanus neptuni – p. 57, pl. 26, figs. 1–6. Hably 1979 [20]: Platanus neptuni – p. 33, pl. 8, fig 5, pl. 9, figs. 1–3, 5, 6, pl. 10, pl. 11, figs. 1, 3, 4. Hably 1980 [20]: Platanus neptuni – p. 300, pl. 2, figs 3–9, pl. 3, pl. 4, pl. 5, figs. 1, 3, pl. 6–10, text-fig. 2. Hably 1982 [21a]: Platanus neptuni – p. 93, pl. 1, figs. 4, 6, 9–11, pl. 2, figs. 1, 2, pl. 5, figs. 1–6, pl. 6, figs. 1–6, pl. 7, figs. 2–4, pl. 8, fig. 1; Debeya hungarica, p. 96, pl. 2, figs. 8, 10, pl. 3, figs. 1–3, 5– 9, pl. 9, figs. 4–5, pl. 10, figs. 1–4, pl. 11, figs. 1–4, pl. 12, fig. 1; Plantae incertae sedis – p. 102, pl. 3, fig. 4, pl. 12, fig. 4. Kovar 1982 [13]: Platanus neptuni – p. 53, pl. 8, figs. 1–10, pl. 9, figs. 7–10, pl. 21, figs. 1–6, pl. 22, fig. 2, pl. 29, figs. 1–6, pl. 36, figs. 1–2. Bu˚zˇek et al. 1982 [8c]: Platanus neptuni – p. 62. Knobloch and Kvacˇek 1983 [12]: Platanus neptuni – p. 99, pl. 4, figs. 4–6. Friis 1985 [5]: Platanus neptuni – p. 29, pl. 4, figs. 8–9, pl. 5, figs. 1–5. Hably 1985a [22b]: Platanus neptuni – p. 145, pl. 17, fig. 6, pl. 18, figs. 1–6, pl. 19, figs. 1–5, pl. 20, figs. 1–4, pl. 21, figs. 1,-3; Laurus princeps – partim, p.143, pl. 14, fig. 3; Laurophyllum pseudoprinceps – p. 144, pl. 17, figs. 2–3. Hably 1985b [20, 21a, d, 22 b]: Platanus neptuni – p. 37, text-figs. 2–4. Mai and Walther 1985 [7a]: Platanus fraxinifolia, p. 55, pl. 13, figs. 1–8, text-figs. 9/3–6, 13/1–2. Mai and Walther 1985 [7b]: Platanus neptuni – p. 54, pl. 12, figs. 23–28. Walther 1985 [7a,]: Platanus fraxinifolia – p. 14, text-figs. 1/6–7. Walther 1985 [7 c, e]: Platanus neptuni – p. 12, text-figs. 1/1–5, 2/1, 3–4. Walther 1985 [7b]: Platanus neptuni – p. 12, text-figs. 2/2, 5. Mihajlovic 1985 [27]: Platanus neptuni – p. 355, pl. 23, fig. 13, pl. 24, figs. 1–3. Cˇernjavska, Palamarev and Petkova 1988 [30b]: Panax longissimum – p. 31; Dewalquea fraxinifolia – p. 31, pl. 3, fig. 4. Hably 1988 [21b]: Platanus neptuni – p. 39, pl. 6, fig. 5, pl. 7, figs. 1–5. Hably 1989 [21c]: Platanus neptuni – p. 87, pl. 3, fig. 3; Platanus fraxinifolia – p. 87, pl. 3, fig. 6, pl. 4, figs. 5–6. Hably 1990 [21d]: Platanus neptuni – p. 15, pl. 13, figs. 1–3, pl. 14, figs. 1–3, pl. 15, 1–3, pl. 16, figs. 1–3, pl. 17, figs. 1–2, pl. 18, fig. 1, text-fig. 53–82.

Z. Kvacˇek and S. R. Manchester: Platanus neptuni complex from the Tertiary of Europe Kvacˇek and Bubı´ k 1990 [15] Platanus neptuni – p. 89, pl. 4, fig. 3, text-fig. 5. Bu˚zˇek et al. 1990 [8b, 9f]: Platanus neptuni, p. 173, 176, text-figs. 5.5, 5.6, 5.12, 5.16, 10.5. Kvaeèk and Hably 1991 [23c]: Platanus neptuni – p. 55, pl. 5, fig. 4. Mai and Walther 1991 [6, 7d, e]: Platanus neptuni – p. 49, pl. 5, figs. 10–14. , pl. 21, figs. 17–19. Ja¨hnichen 1991 [24a, b]: Platanus neptuni – p. 165, pl. 1, figs. 1–2, text-figs. 1–5. Bu˚zˇek, Dvoøa´k, Kvacˇek and Proksˇ 1992 [9h]: Platanus neptuni – p. 125, pl. 3, fig. 1. Sita´r and Kvacˇek 1993 [16a, b, 22a]: Platanus neptuni, Debeya hungarica – pp. 77–78. Hably 1993 [23d] Platanus neptuni – p. 10, pl. 2, fig. 1. Knobloch 1994 [9c]: Platanus neptuni – p. 65, pl. 1, fig. 1. Hably 1994 [21e]: Platanus neptuni – p. 16, pl. 6, figs. 3–6; Platanus fraxinifolia – p. 17, pl. 7, figs. 1–5, pl. 8, figs. 1–4, pl. 9, fig. 4, text-fig. 27. Uzunova 1995 [28]: Platanus neptuni – p. 5, pl. 2, figs. 1–4. Kvacˇek and Walther 1995 [9e]: Platanus neptuni, p. 205, pl. 1, fig. 1. Palamarev and Staneva 1995 [30a]: Platanus neptuni – p. 122, pl. 4, figs. 2–3, 5; Dewalquea fraxinifolia – p. 115, pl. 3, figs. 1, 6–7, 9. Bu˚zˇek, Holy´ and Kvacˇek 1996 [8d]: Platanus neptuni – p. 205, pl. 10, fig. 5, pl. 14, figs. 5–6. Kovar-Eder et al. 1996 [21b]: Platanus neptuni – p. 403, pl. 30.1, fig. 1. Knobloch, Konzalova´ and Kvacˇek 1996 [8a]: Platanus cf. fraxinifolia – p. 55, pl. 18, fig. 8, pl. 21, fig. 5, pl. 22, figs. 1–2, pl. 23, fig. 1, pl. 36, fig. 3, pl. 50, fig. 4, text-fig. 15 a–e. Mihajlovic 1996–1997 [25]: Platanus fraxinifolia – p. 77, pl. 2, figs 1–8; Platanus neptuni – p. 77, pl. 2, fig. 13. Sita´r and Kvacˇek 1997 [22a]: Platanus neptuni – p. 230, pl. 3, figs. 9–14, pl. 7, fig. 5. Mai 1997 [11a]: Platanus neptuni – p. 34, pl. 6, figs. 15–16. Kvacˇek and Walther 1998 [9d]: Platanus neptuni – p. 24, pl. 12, figs. 9–15, text-figs. 13/12–13. Kvacˇek 1998 [9h]: Platanus neptuni – p.131, text-fig. 3.12. Palamarev, Kitanov, Bozukov and Staneva 1999 [29a]: Platanus neptuni – p. 12, pl. 3, fig. 3;

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Myrica lignitum – p.15, pl. 4, fig. 6; Celastrophyllum andromedae (Ung.) Palamarev et al., p. 19, pl. 6, fig. 10; Celastrophyllum serratum – p. 19, pl. 10, fig. 5; Dewalquea fraxinifolia – p. 20, pl. 7, fig. 3, pl. 8, fig. 2, pl. 10, fig. 2. Palamarev, Kitanov and Bozukov 1999 [29b]: Platanus neptuni – p. 32. Walther 1999 [11b]: Platanus neptuni – p. 94, pl. 8, figs. 6–8, text-fig. 16/18. Stro¨bitzer 1999 [12]: Platanus neptuni – p. 98, pl. 2, figs. 1–2, pl. 9, figs. 1–2. Mai 1999 [11b]: Platanus neptuni – p. 22, pl. 7, fig. 19. Hably, Erdei and Kvacˇek 2001 [9a, g]: Platanus neptuni – pp. 14, 17, 28, 37–39, 47, 53, pl. 8, fig. 1, pl. 21, figs. 4–5, pl. 33, figs. 4–5, pl. 34, figs. 3, 5, pl. 38, fig. 2, pl. 51, figs. 3–4, pl. 63, fig. 1. Kvacˇek, Manchester and Guo 2001[7a, 8c, 9a, d, 10a]: Platanus neptuni – p. 453, figs. 7A, C, D, 8B, E, F; Platanus fraxinifolia, p. 453, fig. 7B. Kvacˇek 2002 [9b]: Platanus neptuni – p. 222, pl. 3, fig. 1. Neˇmejc et al. 2003 [8e]: Platanus neptuni – p. 125, pl. 1, fig. 10, pl. 3, fig. 11. Radoo`, Kvacˇek and Walther in press [9e]: Platanus neptuni. Kvacˇek in press [3]: Platanus neptuni. Kvacˇek [17]: Platanus neptuni – (pers. observation Z.K., coll. NHMV No. B 1878.VI.8411). Kvacˇek and Velitzelos (in progress) [31]: Platanus neptuni. This paper [1, 3, 7a, e, 8c, d, 9a, d, e, h, 10a]. We acknowledge private collectors J. Valı´ cˇek and Z. Dvorˇ a´k for donation and loan of rare fossil material. We appreciate exchange of views and other information on Platanus neptuni supplied by our colleagues Harald Walther, Lilla Hably and Dieter H. Mai and have benefited from valuable comments by the reviewers Thomas Denk and Susan Magalloń. We are also thankful for the access to facilities extended by the curators of collections at NM Prague, MMG Dresden, SM.G Frankfurt, USNM Washington and other institutions. The research was financially supported by the Ministry of Education CR (Grant projects No. 205/01/B-GEO/PrF 2000; MSM 113100006) and NSF grants 560260112 and EAR 0174295.

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Addresses of the authors: Prof. Zlatko Kvacˇek Charles University, Faculty of Science, Albertov 6, Praha 2, CZ-128 43, Czech Republic (e-mail: [email protected]); Prof. Steven R. Manchester, Florida Museum of Natural History, University of Florida, Gainesville, Florida, 32611-7800, USA (e-mail: steven@flmnh. ufl.edu)