Revision of Upper Permian Conifer Ortiseia

Revision of Upper Permian Conifer Ortiseia

by Michael Wachtler P. P. Rainerstrasse 11, 39038 Innichen, Italy; E-mail: [email protected]

Abstract

The conifer Ortiseia can be regarded as the typical Alpine Permian plant-genus. It is widespread in the Dolomites and was found there in several species. Since Ortiseia (leonardii) was established by FLORIN in 1964, based on its leave habitus, several other species have been added. After intensive researches in 1984 the Dutch paleobotanist Johanna A. CLEMENT-WESTERHOF published two additional Upper Permian (Lopingian) species - Ortiseia jonkeri and Ortiseia vissheri - and in 2013 Michael WACHTLER described with Ortiseia daberi the first Lower Permian (Cisuralian) conifer of this genus. Based on rich material, partially from new but also well-known sites, it became obvious that a revision was required. In this context emerged Ortiseia zanettii n. sp., another interesting species characterized by its distinctive Araucarian-like pollen cones, till now not recorded in this articulateness in any other Orteisaceae. April 2015. Key words: Coniferophyta, Ortiseia, Permian floras, Dolomites

Division: Coniferophyta Order: Coniferales (TAYLOR, 1981) Family: Ortiseiaceae (CLEMENT-WESTERHOF, 1984)

and seeds) Wachtler Collection, Museum Dolomythos, Innichen.

Genus: Ortiseia (FLORIN, 1964)

Locus typicus

Emended description

Conifer with erect stem and pinnately branched lateral shoot system. Leaves from leathery to awl-shaped and lanceolate. Male cones terminal and solitary, relatively long and slender. Female cones with only one seed sunk deeply in the scale, surrounded by minute sterile protective leaves. Seed scales dropped completely after maturity.

Ortiseia leonardii (FLORIN, 1964) 1964 Ortiseia leonardii, FLORIN, Pl. 1-3 1984 Ortiseia leonardii (Florin), CLEMENT-WESTERHOF Pl. I-XI pp. 122-135

Holotype

Specimen 1/mo Museum Gherdëina, St. Ulrich, South Tyrol

Additional material

CUEC 63, 69, 43, 73 (Twigs and leaves), CUEC 60, 93, 106, 93, 129 , 139, 99 (Pollen cones), CUEC 112, 57, 10, 2, 30, 7 (Female cones), CUEC 26, 37, 34, 36 (Seed scales

Type horizon and age

Late Permian Wuchiapingian (Lopingian) Cuecenes (Seceda)

Stratum typicum

Gröden Formation

Emended diagnosis

Shoots pinnately branched, bearing hardly overlapping leaves. Foliage fleshy, with an acute to obtuse apex, leaf base slightly contracted. Male cones slender, with long bracts as juveniles, mostly losing this characteristic when mature. Female cones bulky to sometimes ellipsoid. Ovuliferous scales with a fair number of sterile leaves surrounding the seed scale. The single nut-like seed centrally attached and embedded. Seed scale falling off entirely after maturity.

Description

Branches: Pinnately spaced out to the sides. The spirally arranged leaves do not or only slightly overlap. CUEC 63 shows well the structure of a Ortiseia leonardii-shoot. Leaves: Ovate to lanceolate with nearly invisible grooves across the entire foliage. Apex acute to obtuse. Leave-base slightly

Michael Wachtler and Thomas Perner: Permian Fossil plants from Europe

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contracted. Single Ortiseia-leaves are normally from 1.0 (CUEC 69) to 1.5 cm (CUEC 43) long and at the base up to 0.5 to 0.8 cm wide. Male cones: Slender, up to 10 cm long (CUEC 106), about 2 cm wide, with an axis bearing numerous spirally arranged and overlapping microsporophylls. Young cones characterised by microsporophylls with extremely long bracts up to 1.5 cm (CUEC 93, 129), adult cones (CUEC 106, 139) partially losing this characteristic. Pollen sacs difficult to establish if they can be regarded as araucaroid, with a fair number of pollen sacs hanging from the outer lower margin towards the main axis. Ovuliferous cones: Usually bulky to round bodied, at least 5 cm long (CUEC 10), and 2 to 2.5 cm wide (CUEC 112, 57, 10, 2, 30, 7). Cone axis holding spirally arranged seed scales. Various minute and elongated sterile leaves cover the coalesced four-lobed seed scale densely on the outer side. Ovule/seed approximately in the middle, up to 1 cm long and 0.5 – 0.7 cm wide (CUEC 26, 34, 36). Seed scales, 1.2–1.5 cm long, 1.0 cm wide (CUEC 37) dropped after maturity as one unit. No distinct and overlapping bract recognizable.

possible to revise in some parts the wellconsidered work of CLEMENT-WESTERHOF. Corrections and adaptations were made on the polliniferous cones in various growth stages, introducing with Ortiseia zanettii the first unambiguous araucaroid microsporophylls and especially the structure of the ovuliferous cones and the composition of the seed scale and seedcomplex. For Ortiseia leonardii WACHTLER concentrated his researches on the locus typicus Seceda/Cuecenes. The exact locality was shown to him by Alfons MORODER, the son of Heinrich MORODER (1896-1974). This indefatigable collector discovered the site on the dangerous slopes of Cuecenes, meaning in the native Ladinian language “red valley”. He handed all the material to Piero LEONARDI (1908-1998), who then passed the fossils without comment to the famous paleobotanist Rudolf FLORIN. Tragically not Moroder but Leonardi was honoured for this “postmaster service” with the name Ortiseia (the hamlet in the Dolomites where the Seceda lies) leonardii.

Emended discussion

Although the genus Ortiseia (leonardii) was described by FLORIN in 1964, based on its leave habitus, it has since then emerged that the characteristic features distinguishing all Permian conifers depend on the construction plan of their cones and even more on the arrangement and composition of the seedscales. Therefore the Dutch researcher Johanna A. CLEMENT-WESTERHOF started intensive studies and published results in 1984 based mainly on the fertile organs and the cuticles, establishing as part of Ortiseia two new Upper Permian species: Ortiseia jonkeri and Ortiseia vissheri (CLEMENT-WESTERHOF, 1984). They were found in other localities in the Dolomites (Bletterbach) and the Vicentinian Alps (Recoaro and Valli del Pasubio). From 2011 to 2013 Michael WACHTLER conducted further studies on Artinskian/Kungurian localities in the Dolomites (Tregiovo), describing first a Lower Permian Ortiseia-species (Ortiseia daberi) and then reworking known Upper Permian material but also new localities from the Dolomites and the Vicentinian mountains. Due to the rich material it was

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Ortiseia leonardii. Reconstructions of single shoots and leaves considering species on the right side (CUEC 69, 43, 73).

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Ortiseia leonardii: Twigs and leaves 1. CUEC 63 Ortiseia leonardii. Branch

2. CUEC 69 Ortiseia leonardii. Apical part of a twig 3. CUEC 43 Ortiseia leonardii. Single leave. 4. CUEC 73 Ortiseia leonardii. Leaves on a main stem


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Ortiseia leonardii: Pollen cones and microsporophylls

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5. CUEC 60 Ortiseia leonardii. Juvenile male cone

6. CUEC 93 Ortiseia leonardii. Mainly adult male cone. Note the long bracts 7. CUEC 106 Ortiseia leonardii. Adult male cone. Note the dimension of the bracts 8. CUEC 93 Ortiseia leonardii. Detail of an isolated microsporophyll 9. CUEC 129 Ortiseia leonardii. Juvenile microsporophylls had longer bracts than the adult one 10. CUEC 139 Ortiseia leonardii. Adult male cone. Apical side 11. CUEC 99 Ortiseia leonardii. Detail of the microsporophylls. Lateral side.

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Ortiseia leonardii: Female cones and macrosporophylls and ovules 12. CUEC 112 Ortiseia leonardii. Young female cone

13. CUEC 112 Ortiseia leonardii. Two megasporophylls, abaxial view. The ovules are hidden behind the sterile leaves. 14. CUEC 57 Ortiseia leonardii. Detail of one juvenile megasporophyll in a cone. The ovule lies mainly invisible in the background 15. CUEC 10 Ortiseia leonardii. Mainly mature female cone 16. CUEC 2 Ortiseia leonardii. Fully grown female cone. The bushy protectve leaves cover the seed 17. CUEC 30 Ortiseia leonardii. Adult female cone attached on a branchlet 18. CUEC 7 Ortiseia leonardii. Mature cone loosing the seed-scales


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Ortiseia leonardii: Seeds

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19. CUEC 37 Ortiseia leonardii. Dropped megasporophyll with hidden ovule, abaxial view (outside) 20. CUEC 26 Ortiseia leonardii. Megasporophyll adaxial view, internal side 21. CUEC 34 Ortiseia leonardii. Seed overview 22. CUEC 36 Ortiseia leonardii. Seed scale evidencing the form of the seed

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Ortiseia. Growing stage from the ovule to the seed a. Juvenile megasporophyll, the ovule in the background, abaxial view (CUEC 112), Adaxial view of the juvenile ovule, covered in a circular pattern by protective leaves, c. Mature megasporophyll, abaxial side (CUEC 11), d. Mature megasporophyll, surrounded by sterile protective leaves, overview (CUEC 34), e. Megasporophyll opening to show the seed PAS 80, f. Mostly open seed (CUEC 262), g. Seed reverse view ULBE 40), h. Seed, obverse view (ULBE 50)

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d e Ortiseia leonardii. Reconstruction. a. Twig CUEC 63), b. Isolated leave (CUEC 43), c. Lateral view of a leave, d. Twig with two female cones, e. Pollen cone (CUEC 93), f. Isolated microsporophyll outer side (CUEC 93).


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Ortiseia jonkeri (CLEMENT WESTERHOF, 1984) 1984 Ortiseia jonkeri CLEMENT-WESTERHOf Pl. XX-XXI pp. 142-160


Late Permian Wuchiapingian (Lopingian)

Emended diagnosis

Cortiana, Valli del Pasubio

Pinnately branched shoots situated in one plane. Leaves awl-shaped, incurved. Male cones slightly bulgy, microsporophylls with a leafy appendix. Female cones bulky to ellipsoid. Ovuliferous scales with a fair number of sterile leaves surrounding the seed scale. Only one nut-like seed centrally attached. Seed scales falling off entirely after maturity.

Stratum typicum

Description

Holotype

Specimen 14108 Laboratory of Palaeobotany and Palynology of the State University of Utrecht The Netherlands.

Locus typicus

Gröden Formation

Additional material

ULBE 01, 23, 80 (Twigs and leaves), ULBE 59, 62, 110, 01, 12, 10 (Pollen cones), ULBE 61, 18, 19, 66, 15 (Female cones), ULBE 35, 40, 50, 116 (Seed scales and seeds). All Wachtler Collection, Museum Dolomythos, Innichen

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Branches: Widely spaced (ULBE 01) and symmetrically spreading out to the sides with slightly imbricate leaves. Leaves: Awl-shaped incurved (ULBE 23, 80) to triangular (ULBE 140) reaching about 1 cm in length and 0.5 – 0.7 in width. Male cones: Fully grown cones reach a length from 4.5 cm (ULBE 59) to 6 cm (ULBE 62) and about 2 to 3 cm in width.

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1. ULBE 01 Ortiseia jonkeri. Twig with slightly incurved needles

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Ortiseia jonkeri: Twigs and leaves

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2. ULBE 140 Ortiseia jonkeri. Branchlet with needles 3. ULBE 23 Ortiseia jonkeri. Branchlet with needles 4. ULBE 80 Ortiseia jonkeri. Branchlet

They are elongated and slender, sometimes bulging in the middle. From the axis (about 3 cm thick) sprout numerous spirally arranged and overlapping microsporophylls (ULBE 12) that usually reach a length of 0.5 cm. The microsporophylls end in a short pointed bract of about 0.5 cm (ULBE 12). If the structure of the pollen sacs is typical araucaroid is difficult to establish. Female cones: From round-bodied, 2.5 cm length and 2 cm in width (ULBE 19) till elongated with a maximum 5 cm length (ULBE 66) and 3 cm width (ULBE 18). Cone axis holding spirally arranged seed scales, bearing on their inner side the ovuliferous seeds. They are composed of various minute and elongated sterile leaves surrounding the seed scale. This coalesced four-lobed structure holds one rounded to slightly elongated ovule/seed in the approximate middle. Seed scales 1.2 – 1.5 cm long, dropped after maturity as a single unit. No visible projecting bract. Seeds about 1 cm long, 0.6 – 0.8 cm wide, surrounded initially by the fleshly scale and protective leaves, then gradually losing them.

Emended discussion

Apart from CLEMENT-WESTERHOF’s locus typicus Cortiana in the Valli del Pasubio, Ortiseia jonkeri was also recovered in the nearby Recoaro hamlet Ulbe. This site is characterized by its richness in male and female cones. The differences from Ortiseia leonardii consist mainly in differences in the male cones, which hold shorter bracts in O. jonkeri (0.5 cm against 1 cm). The aspect of the female cones is nearly identical. More discrepancies can be seen with Ortiseia zanettii from the nearby Ariche locality. There the male cones can be interpreted as typically araucaroid, with pollen sacs hanging from the outer margin, a feature not clearly recognizable in O. jonkeri. Also, the size of the male cones is twice that in the other species, whereas the single microsporophylls in O. zanettii are huge and bulky with a thickness of about 1 cm. In this O. jonkeri has more parental affinities with O. leonardii. The probably fasten mutation from more Walchian to more Araucarian conifers - based particularly on the aspect of the male cones (the females remained nearly equal) - has to be studied in more detail.


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Ortiseia jonkeri: Pollen cones

5. ULBE 140 Ortiseia jonkeri. Isolated needle 6. ULBE 63 Ortiseia jonkeri. Juvenile male cone 7. ULBE 59 Ortiseia jonkeri. Juvenile male cone 8. ULBE 62 Ortiseia jonkeri. Pollen cone evidencing the outer part and the microsporophylls from the inner side. 9. ULBE 07 Ortiseia jonkeri. Semi-destroyed old male cone 10. ULBE 12 Ortiseia jonkeri. Detail of the axis 11. ULBE 10 Ortiseia jonkeri. Microsporophylls from the lateral side

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Ortiseia jonkeri: Female cones

12. ULBE 61 Ortiseia jonkeri. Juvenile female cone attached to the branchlet 13. ULBE 18 Ortiseia jonkeri. Mature female cone with seeds ready to be released 12. ULBE 67 Ortiseia jonkeri. Apical part of a female cone 13. ULBE 15 Ortiseia jonkeri. Female cone evidencing the bushy protective leaves 14. ULBE 66 Ortiseia jonkeri. Fully grown female cone


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Ortiseia jonkeri: Ovules and seeds

15. ULBE 35 Ortiseia jonkeri. Ovule covered by protective leaves 16. ULBE 40 Ortiseia jonkeri. Seed reverse view 17. ULBE 50 Ortiseia jonkeri. Seed obverse view 18. ULBE 116 Ortiseia jonkeri. Colluvium of seeds

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Ortiseia zanettii n. sp. (WACHTLER, 2015)


Holotype

Etymology

PAS 94 Dolomythos Museum, Innichen, Coll. Wachtler

Paratype

PAS 21, 57 (Male cone and microsporophyll), PAS 379 (Female cone).

Locus typicus

Ariche, Valli del Pasubio

Stratum typicum

Gröden Formation

Additional material

PAS 512, 589 (Twigs and leaves), PAS 94, 304, 637, 612, 56, 487, 60 (Pollen cones), PAs 69, 12 (Female cones), PAS 554, 364, 352 (Seed scales and seeds).

Repository

All Wachtler Collection, Museum Dolomythos, Innichen

Late Permian Wuchiapingian (Lopingian) Named after Feruccio Zanetti, the owner of the terrain, who generously allowed the material to be collected.

Diagnosis

Incurved to close fitting, slender leaves. Male cones long and slender, microsporophylls massive with a broad appendix. Pollen sacs easily visible, typically araucaroid, hanging from the apical side of the microsporophyll dorsiventrally to the main axis. Female cones ellipsoid to elongate. Ovuliferous scales with a fair number of sterile leaves covering the outer side of the seed scale. Only one nut-like seed centrally attached. The seed scale falls off entirely after maturity. Seeds rounded to slightly elongated.

Description

Branches: Fine slender leaves covering the main stems (PAS 694).

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1. PAS 363. Ortiseia zanettii n. sp. Slab with two female cones (a = lateral view, b. overview), c. twig, d. single seed


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2. PAS 94 Ortiseia zanettii n. sp. Designed holotype. Branchlet with needles 3. PAS 589 Ortiseia zanettii n. sp. Branchlet with needles 4. PAS 694 Ortiseia zanettii n. sp. Stem axis

Leaves: Slightly awl-shaped to linear, 1 cm long but only 1 mm wide, fitting close to the stem (PAS 512, 589, 94). Male cones: Up to 10 cm long (PAS 51), about 2.5 (PAS 51) to 3 cm wide (PAS 56), elongated, having the same size at the beginning as on the end (PAS 304, 637, 612, 56). Microsporophylls up to 1 cm long and 0.5 cm wide, massive, enlarged with a short broad apex (PAS 219, PAS 133). Tiny hairs sometimes visible. Pollen sacs, massive, close hanging on the lower apical part towards the main axis, probably 4 on each row (PAS 21, 94). Female cones: Round bodied to elongated, 3.5 long, 2 cm wide (PAS 379). Cone axis holding spirally arranged seed scales. These are 1.5 cm long and 0.8 cm wide, covered on the outer side with various minute sterile protective leaves (PAS 554). Seed scale coalesced, four-lobed, holding a 1.0 cm long and 0.5 cm wide rounded ovule/seed approximately in the middle, dropped after maturity as a single unit. No visible bract.

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Discussion

This newly discovered Ortiseia species stands apart from the till now described Ortiseia leonardii, O. vissheri and O. jonkeri due to its distinctive pollen cones. They are much more massive than those of Ortiseia leonardii or O. jonkeri, and are the only ones where it is possible to recognise the typical blueprint of all extant Araucarias: The pollen sacs hang from the apical lower side dorsiventrally to the main axis. Strangely, in no other Permian Ortiseia or Walchiaspecies can this feature be recognised. In this they resemble the male cones of the Triassic Voltzia. Therefore, the discovery of Ortiseia zanettii is important to understand the evolving line of Araucariaceae. Although in almost all paleobotanical literature the visual aspect of Paleozoic-Mesozoic pollen cones is defined as uniform, it can be established that it is not so. The huge Permian Ortiseia-Araucaria cones stand in stark contrast to the small Majonica-Picea pollen

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Ortiseia zanettii: Female cones

5. PAS 379 Ortiseia zanettii n. sp. Paratype. Fully grown female cone 6. PAS 69 Ortiseia zanettii n. sp. Broken female cone 7. PAS 12 Ortiseia zanettii n. sp. Mature female cone, ready to release the seed scales 8. PAS 628 Ortiseia zanettii n. sp. Young female cone


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Ortiseia zanettii n. sp.: Ovules and seeds 9 - 11. Ovules outside (PAS 554, 364, 352) 12 - 14. Ovules overhead view and inner face (PAS 6, 89, 80) 15 - 16. Seeds obverse and reverse view (PAS 77, 408)

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Ortiseia zanettii n. sp.: Cone ecology

17. PAS 56 Ortiseia zanettii n. sp. Deposit of three adult male cones 18. PAS 345 Ortiseia zanettii n. sp. Deposit of a male (a) and female (b) Ortiseia-cone, a Ortiseia seed-scale (c) and a Majonica seed-scale (d) with winged seeds 19. PAS 695 Ortiseia zanettii n. sp. Deposit of a young female and a pollen cone


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20. PAS 94 Ortiseia zanettii. Designed holotype. Juvenile male cone attached on a twig 21. PAS 304 Ortiseia zanettii. Entire juvenile pollen cone 22. PAS 637 Ortiseia zanettii. Pollen cone with outer bracts 23. PAS 51 Ortiseia zanettii. Adult male cone 24. PAS 60 Ortiseia zanettii. Semi destroyed male cone 25. PAS 57 Ortiseia zanettii. Old pollen cone without pollen sacs

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Ortiseia zanettii: Pollen cones

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26. PAS 94 Ortiseia zanettii. Microsporophyll with hanging pollen sacs 27. PAS 219 Ortiseia zanettii. Paratype. Microsporophyll evidencing well the hanging pollen sacs

Ortiseia zanettii n. sp. Reconstruction of single microsporophylls from diverse sides and the whole pollen cone.


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cones and, in almost the same manner, to the male cones of Valentinia-Pinus. There are fewer differences between Ortiseia zanettii and the other Ortiseia-species in the look of their ovuliferous organs. Their coalesced seed-scale concept with various sterile tapered minute leaves, which serve to cover the seed scale, survived in slightly modified form till today’s Araucarias. But this can serve as a distinctive character in contrast to other Permo-Triassic conifers like the winged seeded Wachtlerina, Majonica or the Pinus-like Valentinia. Surprisingly, and in net contrast to the abundant branches and rich leaves in Seceda-Cuecenes locality where Ortiseia leonardii is the dominant conifer, in the locus typicus for Ortiseia zanettii - the Contrada Ariche (Valli del Pasubio) - well preserved leaves and branches are rare, while both female and male cones are abundant and also well maintained. It is a mystery why they are missing whilst others, like fragile Cycad-leaves, were found in quite good quantities. Probably all different Ortiseia species of the Southern Alps can be attributed to some slight age-differences, whereby it could not be established exactly which of the SouthAlpine Upper Permian localities are younger or older. From the much more evolved and, in contrast to the other sites, species rich Ariche-locality it seems that it must be the youngest, possessing a lot of Early Triassic plant affinities like abundant cycadophyte Nilssonia or Bjuvia-species as well as sphenophyta like Equisetites or some enigmatic proto-angiospermous plants. Returning to the exhausting work and according to CLEMENT-WESTERHOF the building concept of Permian conifer ovulatescales is not so easy to understand. It was influenced for decades by FLORIN’s Walchia dogma, meaning that Permian conifer seed scales evolved as inflorescences and formed the basis for the development of all conifers existing today through the continuous merging of sterile bracts and fertile seed scales (FLORIN 1938-1945). Unfortunately, this theory was based on just a few isolated and often poorly preserved pieces of evidence collected from the Thuringian Forest and, to make matters worse, this mixed group of specimens had also been taken from different strata and time periods.

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Since the discovery of CarboniferousPermian Perneria (WACHTLER, 2013), we know that the conifer seed scale is a complex single y to two or maximum four y-dichotomising fertile leaf, mainly developing after to Zimmermann’s sustainable “emergences and telome theory“ from Devonian plants like Psilophytes (ZIMMERMANN, 1954) just a hundred million years before. Sterile as well fertile leaves evolved according to this concept by dichotomous branching of the axis. The sporangia occupy the apical parts of the leaves. The Ortiseia ovulate seed scale is composed of three different units: a bract, in this case consisting of an agglomeration of many sterile minute leaves on the outer side of the seed scale, as seen well in immature female cones (CUEC 112, 57); a single scale formed as a three times segmented leaf (CUEC 26, PAS 89, 80), holding in the middle a submerged seed. With this concept it is also easy to understand the evolutionary steps of the Auracaria family having the same blueprint. CLEMENT-WESTERHOF, J., 1984. Aspects of Permian Palaeobotany and Palynology. IV. The conifer Ortiseia from the Val Gardena Formation of the Dolomites and the Vicentinian Alps (Italy) with special reference to a revised concept of the Walchiaceae (GOEPPERT) SCHIMPER. - Rev. Palaeobot. Palynol., n. 41, pp. 51166 FLORIN, F. 1938-1945. Die Koniferen des Oberkarbons und des Unteren Perms. - Palaeontographica, 65, Abt. B, 1-729, 65 Fig, 186 Taf, Stuttgart. FLORIN, R., 1964. Über Ortiseia leonardii n. gen et sp., eine Konifere aus den Grödener Schichten im Alto Adige (Südtirol). - Mem. Geopaleont. Univ. Ferrara, 1(1), pp. 3-11, n. 41, pp. 51-166. WACHTLER M. 2013. Early Permian) Flora from Tregiovo-Le Fraine in the Val di Non (Trentino, Northern Italy) - Additional and revised edition WACHTLER M. 2013. Protoconiferophyta from the Carboniferous-Permian (Kasimovian/Gzhelian) Niederhausen Flora (Rheinland-Pfalz, Germany) in Perner & Wachtler: Permian fossil plants in Europe and their evolution, Dolomythos and Oregon Institute of Geological Research, Portland ZIMMERMANN W., 1954. Ueber die mikrophyllen Psilophyten, ihre Entstehung und Bedeutung fuer die Stammesgeschichte. Palaeontologische Zeitschrift 28(12): 56-66

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