Ettingshausenia Stiehler, 1857

Halamski, Adam T., Kvaček, Jiří, Svobodová, Marcela, Durska, Ewa & Heřmanová, Zuzana, 2020, Late Cretaceous mega-, meso-, and microfloras from Lower Silesia, Acta Palaeontologica Polonica 65 (4), pp. 811-878 : 834-840

Genus Ettingshausenia Stiehler, 1857

Type: Ettingshausenia cuneifolia ( Bronn, 1837) Stiehler, 1857 ≡ Credneria cuneifolia Bronn, 1837 . Cenomanian, Upper Cretaceous; Nied- erschöna, Saxony.

Remarks.— Ettingshausenia may be understood either broadly ( Maslova et al. 2005; Herman and Kvaček 2010) or narrowly ( Golovneva 2011). Following arguments given by Halamski and Kvaček (2015: 119) the former intepretation is used herein.

Ettingshausenia cf. superstes ( Velenovský, 1882) Kvaček and Halamski in Halamski and Kvaček, 2015

Fig. 13A–C, E .

Material.—Czaple, Assemblage 2, Coniacian: MB.Pb.2008/ 0369. Wartowice, Assemblage 2, Coniacian: MB.Pb.2008/ 0332, 0333, 0334. Zbylutów, Assemblage 2, Coniacian: MB.Pb.2008/0336.2.

Description.—The available material consists of fragments of medium-sized leaves with weakly developed lobes.

Lamina probably notophyll, in the most complete specimen ( Fig. 13B 2) observed length about 6.7 cm and estimated laminar length-to-width ratio about 1.5. Base sharply cuneate ( Fig. 13B ); lateral lobes weakly developed, incised to about one tenth of the blade ( Fig. 13C, E ); a single observed large lobe ( Fig. 13E ) interpreted as the terminal one. Venation palmately pinnate brochidodromous to camptodromous with strong basal veins emerging from leaf base. Tertiary veins percurrent ( Fig. 13B 1 ).

Remarks.—This leaf type is similar to Ettingshausenia superstes , from the Coniacian of the Bohemian Cretaceous Basin ( Velenovský 1882: 15–16 [8–9], pl. 4 [2]: 7–9; Halamski and Kvaček 2015: 311–312, fig. 6I–K), but differs in having small lobes which are not preserved in the holotype. Ettingshausenia onomasta ( Bayer, 1896) Kvaček and Halamski in Halamski and Kvaček, 2015, from the Coniacian of Idzików ( Halamski and Kvaček 2015) differs from the studied material in having well-developed lobes, which are acute-attenuate. Ettingshausenia gruenbachiana Herman and Kvaček, 2010 , from the Campanian of Austria and Ettingshausenia senonensis ( Knobloch, 1964) Kvaček and Váchová, 2006 , are entire-margined, yet both differ from the studied material in having well-pronounced lobes.

Ettingshausenia sp. 1

Figs. 13D, F, G , 15F, G .

Material.— Ulina, Assemblage 6, lower–middle Santonian: MGUWr 5616p.b (coll. Goeppert), MB.Pb.2008/0264, 0338.

Description.—The available material consists of subcomplete to incomplete relatively large, trilobed leaves.

Lamina notophyll to mesophyll, maximal observed length about 9 cm, laminar length-to-width ratio about 1.1 ( Fig. 13G ) to 1.6 ( Fig. 13D ). Base cuneate. Lobes incised to one seventh to one sixth of the length. Venation palmately pinnate with robust basal veins emerging from the leaf base. Secondaries excurrent, departing at 30° ( Fig. 13D ) to 60° ( Fig. 13G ). Percurrent tertiary veins poorly preserved.

Remarks.— Ettingshausenia sp. 1 differs from Ettingshausenia cf. superstes in displaying stronger lobes. Similarities with Ettingshausenia onomasta from the Coniacian of Idzików ( Halamski and Kvaček 2015) should be noted; however, the fragmentary character of the specimen, including in particular a poorly preserved leaf margin, precludes further comparison, so open nomenclature is used here. Variability of leaves classified here as Ettingshausenia sp. 1 is significant; it may correspond to polymorphism within a single tree, to a single variable species that cannot be further subdivided, to a single species with geographical races or ecotypes ( Sell and Murrell 2018: xi), or finally to several biological species.

The specimen MGUWr 5616p ( Fig. 13F ) shows the co-occurrence of Ettingshausenia sp. 1 (5616p.b, lower leaf) and Dryophyllum westerhausianum (upper leaf, 5616p.a).

Genus Platanites Forbes, 1851

Type: Platanites hebridicus Forbes, 1851 ; Paleocene , Isle of Mull, Inner Hebrides, Scotland, UK .

Remarks.— Platanites hebridicus is a trifoliolate platanoid leaf (see Boulter and Kvaček 1989 for a detailed description). Other species, described in much detail, are the similarly trifoliolate P. marginatus ( Lesquereux, 1873) Johnson, 1996 , from the Upper Cretaceous and lower Tertiary of South and North Dakotas, Montana, Wyoming, Colorado, Utah, and Arizona ( Johnson 1996) and P. raynoldsii ( Newberry, 1868) Manchester, 2014, from the Paleocene of North America Manchester 2014). Platanites resembles Ettingshausenia and extant Platanus , but has compound leaves. However, discriminating between compound and simple leaves can be difficult in incomplete fossils. Fragmentary specimens of Platanites can be distinguished by basal secondaries connecting between the diverging primary veins displaying a chevron pattern. Such a pattern is usually absent in Ettingshausenia and even if it does occur, the chevrons are blunt, never sharp. The poorly preserved available material does not offer any traces of the trifoliolate condition of the original leaves, which anyway is seldom preserved in the fossil record and not necessarily constant in the living plant Manchester 2014: 166). Erlingdorfia Johnson, 1996, is another trifoliolate platanoid distinguished by strongly trilobate median leaflets and bilobate lateral leaflets.

As far as it could be checked, this is the oldest record of the genus Platanites .

Platanites willigeri Halamski and Kvaček sp. nov. Figs. 14, 15B, 16A, B.

Etymology: In honour of Gustav Williger (1856–1937; for biographic notes see Springer 1914: 114–117, Anonymous 1937), in recognition of his contribution to the study of the Cretaceous of Lower Silesia Williger 1882).

Type material: Holotype: MB.Pb.2008/0335,basal part of a median leaf- let; designated herein (Fig. 14B) . Paratype 1: MB.Pb.2008/0328, incomplete median leaflet (about two thirds of the blade, with margin, lacking base and apex) (Figs. 14A, 16A) . Paratype 2: MB.Pb.2018/0054.1, a complete lateral leaflet (Fig. 14C). Both paratypes from the same horizon and probably the same locality .

Type locality: Probably Luisenhain, temporary excavations in the first half of the 20 th century. In the collection the type material is labelled “Bunzlau”, but due to recent mislabelling of some specimens (see Appendix 1, entry on Luisenhain) it is unfortunately impossible to be certain about the original locality.

Type horizon: Czerna Formation, most probably lower or middle Santonian.

Material.— Luisenhain, Assemblage 8, lower–middle Santonian: MB. Pb. 2008/0356 (coll. W. Zimmer 1913), MB. Pb. 2018/0056 (coll. W. Zimmer 1912); Assemblage 8, lower– middle Santonian: MB. Pb. 2008/0321, 0329 (counterpart of the paratype 1); MB.Pb.2018/0046, 0048, 0050, 0051 .

Diagnosis.—Leaf interpreted as trifoliolate ( Fig. 15D ); terminal leaflet ovate, base cuneate; margin serrate, teeth rarely spaced; venation palmately pinnate, palinactinodro- mous; midvein stout, two basal veins emerging from leaf base, secondary veins forming chevron pattern by intercon- nected secondary veins between adjacent primaries in the lower part of the lamina. Presumed lateral leaflets unlobed with pinnate venation.

Description.—The available material consists of isolated, most often incomplete laminae, interpreted as terminal (median) leaflets and in a single case a lateral leaflet belonging to a compound trifoliolate leaf, petiolate and with petiolulate leaflets.

Terminal leaflet: blade attachment marginal, laminar size probably microphyll to notophyll; laminar length-to-width ratio probably ca. 1.5–1.6; laminar shape ovate, unlobed, medially asymmetric.Base symmetrical, cuneate (Fig.14B; concave sensu Ellis et al. 2009). Apex not preserved.

Venation of the median leaflet approximately pinnate with mostly a single primary vein (Fig. 14A), but sometimes the first secondary vein being nearly as thick as the midvein (Fig. 16A) resulting in a quasi-palmate pattern. Compound agrophic veins present. Major secondaries craspedodromous, seldom semicraspedodromous, attachment decurrent, angle approximately uniform, 40–50°, spacing apparently decreasing distally. Marginal veins probably absent. Intersecondaries mostly absent (see Remarks). Tertiaries percurrent, mostly convex, relatively sparsely spaced, angles between them and the midvein mostly acute.

Margin of the median leaflet toothed, tooth spacing irregular. Teeth 1–2 per cm, varying in size, but forming not two distinct size classes; most often supplied by a single principal vein. Sinus shape rounded. Proximal flank weakly convex, distal flank straight to concave.

Presumed lateral leaflet (Fig. 14C): lamina lanceolate in shape, laminar length-to-width ratio ca. 1.7; apex shape rounded, probably retuse; teeth regular, blunt; venation pinnate.

Fig. 14. Platanoid angiosperm Platanites willigeri Halamski and Kvaček , → sp. nov. probably from Luisenhain, North Sudetic Basin , Lower Silesia, Poland, Assemblage 8 (lower–middle Santonian). A. Incomplete median leaflet MB.Pb.2008/0328, paratype (see also Fig. 16A) . B. Basal part of a median leaflet MB.Pb.2008/0335, holotype . C. Complete lateral leaflet MB.Pb.2018/0054.1, paratype .

Remarks.—Platanoid affinities of the described leaves are undeniable on account of the venation pattern and tooth characters. Its architecture and shape show notable similarities to those of Platanites raynoldsii and Platanites marginatus of North America. From those leaves P. willigeri differs in having a less decurrent base. From Platanites hebridicus it differs in having more pronounced lobes and in the presence of lateral leaflets. However, the preservation of our material is fragmentary and no specimen shows the physical connection of leaflets.

Similarities of the presumed median leaflet to Platimeliphyllum palanense (Budantsev, 1983) Maslova, 2002, from the upper Paleocene–lower Eocene of Kamchatka ( Maslova 2002: text-fig. 1; see also Maslova 2008: 1399– 1400) should be noted. However, the generic diagnosis of Platimeliphyllum Maslova, 2002, includes cuticular features (see also Kodrul and Maslova 2007), so this name cannot be used for the material from the North Sudetic Basin. Another similar leaf is Protophyllum zaissanicum Romanova, 1963 , from the uppermost Cretaceous (to lower Paleogene?) of Kyin-Kerish Hill in Kazakhstan ( Romanova 1963; see also Kutuzkina 1974: 149, pl. 72: 1); it differs in more closely spaced leaf margin teeth.

Further leaves showing some characters in common with the laminae interpreted herein as median leaflets of a trifoliolate leaf include Late Cretaceous–Paleogene represen-

Fig. 16. Santonian platanophylls and laurophylls from the North Sudetic Basin, Lower Silesia, Poland. A, B. Platanites willigeri Halamski and Kvaček , → sp. nov. probably Luisenhain, Assemblage 8 (Santonian). A. Enlargement of a part of the paratype MB.Pb.2008/0328 (see Fig. 14A) showing teeth and venation. B. Incomplete leaf MB.Pb.2018/0050 showing venation. C, E. Platanites sp. 1 , Nowe Jaroszowice, Assemblage 7 (Santonian). C. Incomplete presumed median leaflet MB.Pb.2008/344, general view (C 1), enlargements of the base and of the margin and venation under different lightning (C 2, C 3).

E. Incomplete median leaflet MB.Pb.2008/268. D. Laurophyllum sp. , leaf MB.Pb.2008/263, Ulina, Assemblage 6 (Santonian).

tatives of Platimelis Golovneva, 1994 that have relatively densely spaced small, symmetric teeth and cordate bases Golovneva 1994; Moiseeva 2012). Leaves of Evaphyllum Maslova, 2003 (type and only known species E. kamchaticum Maslova,2003 ,fromtheupperPaleocene–lowerEocene of Kamchatka ( Russia); see also Maslova 2010: 1448–1449) are elliptic in shape. Leaves reported from the Coniacian to Campanian Valižgenian taphoflora of Kamchatka by Budancev (1983) as Platanus crednerioides Budancev, 1983 Budancev 1983 : pl. 22: 4), and Pseudoprotophyllum boreale ( Dawson, 1883) Hollick, 1930 ( Budancev 1983: pl. 23: 2), differ from our species, the former in having larger teeth, the latter in having agrophic veins forming a more complex system. Both Pseudoprotophyllum Hollick, 1930 and Pseudoaspidiophyllum Hollick, 1930 , from the Creatceous of Alaska have peltate bases. In Viburniphyllum de Saporta, 1868 , and Grewiopsis de Saporta, 1868 , secondaries are more branched.

Platanophyllum Fontaine, 1889 was used by Arens and Allen (2014: 188) to accommodate simple, unlobed platanoid leaves found in the basal part of the Hell Creek Formation Maastrichtian) in Montana ( USA). Such a treatment seems inappropriate given the fragmentary character of the type material of Platanophyllum ( P. crassinerve Fontaine, 1889 ; Potomac Group, Lower Cretaceous, Virginia, USA).

Stratigraphic and geographic range.—Assemblage 8 lower–middle Santonian) at Luisenhain, Poland. Several specimens in MB are labelled “Bunzlau”, but this locality does not appear on the original labels (see Appendix 1, Luisenhain); all material of this species possessing original labels is from Luisenhain. It is uncertain whether P. willigeri is present at Bolesławiec, Poland.

Platanites sp. 1

Figs. 15D , 16C, E.

Material.—Nowe Jaroszowice, Assemblage 7, lower–mid- dle Santonian: MB.Pb.2008/0268, 0344.

Description.—Leaf interpreted as trifoliolate, but in the available material only simple blades (presumably corresponding to median leaflets) are preserved. Blade attachment marginal, petiol(ul)e very long. Venation palmately pinnate. Secondaries connecting primaries in a way similar to a chevron pattern, with the angle of the secondaries being very wide (over 120°).

Remarks.—Interpretation of this fragmentary material is tentative. The similarity of the shape and architecture of the described phyllomes to those of the central leaflets of Platanites marginatus and P. hebridicus is striking. The long petiol(ul)e is also consistent with such an interpretation. However, it cannot be excluded that all specimens are distorted taphonomically in a similar way, or, in other words, they are lacking analogous parts, and that the original shape of the blade was different.

Viburnum vitifolium de Saporta and Marion, 1878 , was described on the basis of a single specimen from Gelinden (de Saporta and Marion 1878: 73–75, pl. 12: 1) which is early Paleocene (Danian) in age (Kvaček 2010), and is also similar to Platanites sp. 1 described herein; differences include the margin, which is entire in our material and dentate in the Paleocene species.

Comparable leaves were described by Wolfe and Wehr (1987) from the Eocene of Republic (Washington state, USA) as Macginitiea gracilis ( Lesquereux, 1872) Wolfe and Wehr, 1987 . They have a similar arrangement of the basal veins, but are distinguished from the studied material by a sharp characteristic chevron pattern formed by inter- connected secondaries ( Wolfe and Wehr 1987; Manchester 2014) which is absent in the described material (compare Fig. 16C 3 and Manchester 1986: fig. 5A–C). Members of Recent Diploclisia Miers, 1851 ( Menispermaceae ) are similar in shape and in venation; the main veins, however, are craspedodromous ( Yu and Chen 1991: pl. 22: 5, 13).

Poorly preserved MB.Pb.2008/0340, 0341, and s.n. from the same outcrop might belong here.

Figures

Fig. 13. Representatives of the platanoid Ettingshausenia from the North Sudetic Basin, Lower Silesia, Poland. A–C, E. Ettingshausenia cf. superstes (Velenovský, 1882) Kvaček and Halamski in Halamski and Kvaček, 2015, Assemblage 2 (Coniacian). A. Leaf fragment MB.Pb.2008/336, Zbylutów. B. Incomplete leaf MB.Pb.2008/333b, Wartowice; general view (B2), enlargement of the apical part (B1). C. Leaf fragment MB.Pb.2008/334, Zbylutów. E. Leaf fragment MB.Pb.2008/332, Zbylutów. D, F, G. Ettingshausenia sp. 1, Ulina,Assemblage 6 (Santonian?). D. Subcomplete leaf MB.Pb.2008/264. F. Incomplete leaf MGUWr 5616p. G. Incomplete leaf MB.Pb.2008/338.

Fig. 1. Geographic, palaeogeographic, and geologic context of the studied flora. A. Late Cretaceous palaeogeography of Central and Northern Europe after Ron Blakey from Csiki-Sava et al. 2015, modified after data in Chatziemmanouil 1982; Surlyk in Voigt et al. 2008; Halamski 2013; Halamski et al. 2016). The red asterisk shows the presumed position of the studied flora. ESI, East Sudetic Island; WSI, West Sudetic Island. B. Geographical location of the study area in Central Europe. C. Bedrock geological map of northern Silesia (pre-Cenozoic redrawn, simplified and partly corrected after Kozdrój et al. 2001; Cenozoic simplified after Sawicki 1966, 1995) with approximate locations of early, early middle, and late Coniacian palaeoshores (after Leszczyński 2018: fig.40A, B, D).The studied localities (seeAppendix 1) are shown by asterisks, black for those whose location is known in detail, white for those for which a precise location cannot be given.

Fig. 15. Semi-schematic reconstructions of the presumed main forest-forming angiosperms in the Coniacian–Santonian of Lower Silesia, Poland. A. Dewalquea aff. gelindenensis de Saporta and Marion, 1873 (compare Fig. 12A, B). B. Platanites sp. 1 (compare Fig. 16C; lateral leaflets hypothetical, supposed similar to those of Platanites marginatus, see Johnson 1996: fig. 86). C. Dryophyllum westerhausianum (Richter, 1904) Halamski and Kvaček comb. nov. (compare Fig. 8C, E). D. Platanites willigeri Halamski and Kvaček, sp. nov. (compare Fig. 14A, C). E. Dewalquea insignis Hosius and von der Marck, 1880 (compare Fig. 10F). F, G. Ettingshausenia sp. 1 (compare Fig. 13G, D). Not to scale. Drawings by Bogusław Waksmundzki.