Leguminosites hradekensis (E. KNOBLOCH et KVAČEK) KVAČEK et TEODORIDIS 2012

Dedicated in memory of the late FrantiŠek Holý (1935 - 1984), an eminent Czech palaeobotanist, Holý, František, Kvaček, Zlatko & Teodoridis, Vasilis, 2012, A Review Of The Early Miocene Mastixioid Flora Of The Kristina Mine At Hrádek Nad Nisou In North Bohemia (The Czech Republic), Acta Musei Nationalis Pragae Series B 68 (3 - 4), pp. 53-118 : 71-72

Leguminosites hradekensis (E. KNOBLOCH et KVAČEK) KVAČEK et TEODORIDIS , comb. n.

Pl. 2, figs 14-19, pl. 12, figs 4-6

1976

Magnolia hradekensis E. KNOBLOCH et KVAČEK , p. 18, pl. 4, figs 15-19, pl. 5, figs 1-7, pl. 9, figs 1, 9, pl. 15, fig. 14, text-fig. 5 (basionym) (Wackersdorf).

Detached fragmentary leaflets, lamina probably ovate to elliptic, 8–20 mm long and 5–13 mm wide, base and apex not preserved, margin entire, venation brochidodromous, midrib strong, moderate, straight, secondary veins numerous, thinner, straight to slightly curved, looping, alternate to opposite, originating at almost acute angles, intersecondaries obvious, thinner, parallel, tertiary veins alternate percurrent, curved to sinuous, venation of higher orders regular polygonal reticulate, areolation well developed, areoles 3- or 4-sided, veinlets one branched to dichotomous branching. Adaxial epidermis medium cutinized, smooth, non-modified cells polygonal to lobate, 20–25 or up to 35 µm in diameter, anticlinal walls almost straight to strongly wavy, cells narrow elongate over veins, with cylindrical basal cells of serial trichomes 15 µm in diameter, abaxial epidermis delicately cutinized reflecting polygonal to lobate non-modified cells with curved to strongly wavy anticlinal walls, stomata amphicycloparacytic, subsidiary cells sometimes repeatedly subdivided radially, outer circle of subsidiary cells composed of larger cells not much different from the non-modified cells, pairs of slender guard cells 13–16 µm wide and 23–26 µm long, mother cells of undeveloped stomata often present. Cylindrical bases of uniserial trichomes 15–20 µm in diameter scattered among stomata, more densely in some leaf forms, complete trichomes up to 180 µm long rarely preserved with thin-walled apical cell.

D i s c u s s i o n: The record from the Wackersdorf locality is much more completely documented than the first specimens recovered in the Kristina Mine (Kvaček 1966). Therefore the material from Wackersdorf was preferably published ( Knobloch and Kvaček 1976) and the most complete specimens chosen as the type of this foliage species, although in the manuscript (Kvaček 1966) the Kristina Mine was intended to be the type locality. The affinity to the Magnoliaceae is now rejected because of the lack of mesophyllous oil cells and the peculiar stomatal type (see Kvaček et al. 2011). More similar in respect of leaf morphology (short petiolule) as well as anatomy are certainly legume leaflets (for comparison see legume foliage described from the Eocene of North America by Herendeen 1992, p. 137, figs 270-276).

M a t e r i a l: Leaflet compressions isolated on slides, G 9172-9175 (KR 21, 43a, b, 410).

Betulaceae GRAY

Alnus julianiformis (STERNBERG) KVAČEK et HOLÝ Pl. 2, figs 20-21, pl. 12, figs 7-8

1823

1845 1974

Phyllites julianiformis STERNBERG , p. 37, 39, pl. 36, fig. 2 (Bílina).

Fagus feroniae UNGER , p. 106, pl. 28, figs 3-4 (Bílina). Alnus julianiformis (STERNBERG) KVAČEK et HOLÝ , p. 368, pls 1-3, pl. 4, fig. 1, text-fig. 1 (Bílina, Břeštany).

Incomplete leaves and fragments, lamina widely elliptic to obovate, 6–32 mm long and 12–25 mm wide, apex not preserved, probably obtuse or acute, base cuneate with fragmentary petiole, margin irregularly simple serrate, teeth acute, sinus angular, venation craspedodromous/semicraspedodromous, midrib straight, strong, secondaries alternate, thinner, regularly spaced, originating at an angle of 30–40°, curved, looping, tertiary veins alternate percurrent, straight or curved, venation of higher orders regular polygonal reticulate, areolation well developed, 4-sided. Adaxial epidermis medium cutinized, smooth, showing outlines of non-modified cells 12–20 µm in diameter with almost straight anticlines, abaxial epidermis thinly cutinized, showing non-modified cells with shallowly wavy anticlines, stomata anomocytic, elliptic, guard cells ca. 25 µm long and 20 µm wide, scattered four-celled trichome bases ca. 28 µm in diameter.

D i s c u s s i o n: In gross morphology and epidermal anatomy the leaf remains described above correspond to A. julianiformis from the Bílina area ( Kvaček and Holý 1974) and Wackersdorf ( Knobloch and Kvaček 1976). A. trabeculosa HAND. - MAZZ. of East Asia has been suggested as the living relative ( Kvaček and Holý 1974).

M a t e r i a l: Leaf compressions on slides, G 9176- 9183 (KR 143, 144, 147, 151, 194a, b, 224, 225, 248).

Fagaceae DUMORT.

Fagus L. Fagus deucalionis UNGER plexus

Pl. 2, figs 22-27, pl. 3, figs 6-7, pl. 6, fig. 20, pl. 7, fig. 1, pl. 12, figs 9-10

1847

1906

1933

1989

1989

1991

Fagus deucalionis UNGER , p. 101, pl. 27, figs 1-6 (Počerny, in German Putschirn).

Fagus ferruginea AIT. forma miocenica MENZEL , p. 48, pl. 3, figs 4-5, 10-12 (Rauno).

Fagus cf. ferruginea AIT. ; Menzel in Gothan and Sapper, p. 15, pl. 3, fig. 8 (Klettwitz).

Fagus attenuata GÖPPERT ; Kvaček and Walther, p. 214, text-fig. 1f (Hrádek/N., Kristina Mine).

Fagus decurrens C. et E.M. REID; Kvaček and Walther, p. 214, text-fig. 1b-c (Hrádek/N., Kristina Mine).

Fagus menzelii KVAČEK et WALTHER , p. 485, pls 7-8, text-figs 9-10 (Kausche).

Incomplete leaves and fragments, lamina elliptic to widely elliptic to ovate, 12–28 mm long and 8–25mm wide, apex not preserved, probably acute to shortly attenuate, base cuneate with petiole up to 8 mm long, margin partly entire, at places regularly widely simple serrate, teeth acute, rarely blunt, sinus angular, venation simply pseudocraspedodromous, midrib straight, strong, secondaries alternate to opposite, thinner, regularly spaced, straight, originating at an angle of 30 to 40°, curved just before entering the marginal tooth, tertiary veins alternate percurrent, straight or curved, often forked, venation of higher orders regular polygonal reticulate, areolation well developed, 4-sided. Adaxial and abaxial cuticles hardly discernible on macerated samples, more visible directly on the sub-macerated leaf compressions. Cupules broadly ovate to broadly acute, 8–21 mm long, 6–13.3 mm wide, on apices often broken, (probable original length 8–21 mm), valves ovoid to broadly lanceolate, on base rounded, concentrically striate, on surface dense decurrent bases of spines present, stalk conical, short. Fruits elongate ovoid, 9–11 mm long, 4.5– 5 mm wide, triangular, apically narrowing into pointed apex, edges sharp to winged, bluntly rounded on base ( Holý 1975, pp. 32-34, pl. 5, figs 15-17, as Fagus decurrens ).

D i s c u s s i o n: Fagus leaf fossils of the European Early-Middle Miocene were usually assigned to F. attenuata GÖPPERT ( Kvaček and Walther 1989) but the revision by Walther and Zastawniak (1991) corrected the affinity of the type specimen to Alnus (see Walther in Kvaček and Walther 1991, p. 488). Later the leaf fossils of this form have been assigned either to Fagus silesiaca WALTHER et ZASTAWNIAK (1991) based on the Late Miocene material of Sośnica or F. menzelii KVAČEK et WALTHER (1991) based on the Middle Miocene material from Germany (Kausche and other sites). Walther (in Kvaček and Walther 1991, p. 488) confirmed that differences between the two entities are negligible. Associated cupules are also very similar, usually assigned to a single fossil species ( Holý 1975, Kvaček and Walther 1989 as Fagus decurrens C. et E. M. REID, Denk and Meller 2001, Teodoridis 2003). We include the two fossil species based on foliage, which were thriving under quite different environmental conditions into a single species plexus and, contrary to the recent treatment of fossil beach foliage ( Denk 2004), we consider them and other fossil morphotypes of beech in Europe as infraspecific variation. As shown by Denk and Meller (2001) the morphology of fossil cupules of beech conforms to a single species throughout Europe and therefore we suggest assigning the populations of the Kristina Mine and others connected with mastixioid floras (including the Late Oligocene site Počerny) into an informal species plexus Fagus deucalionis .

M a t e r i a l: Leaf compressions on slides, G 9184- 9194, partly missing (KR 5, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 23, 25, 26, 105, 106, 107, 108, 109, 110, 111, 112, 113, 114, 121, 122, 123a, b), numerous cupules, several fruits, G 8821, 8822, Gs 232.

Trigonobalanopsis KVAČEK et WALTHER