Trigonobalanopsis rhamnoides (ROSSMÄSSLER) KVAČEK et WALTHER

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 : 72-75

Trigonobalanopsis rhamnoides (ROSSMÄSSLER) KVAČEK et WALTHER plexus

Pl. 3, figs 1-4, pl. 7, figs 2-3, pl. 12, figs 11-12

1844 Phyllites rhamnoides ROSSMÄSSLER , p. 35, pl. 8, figs 30-31 (Altsattel, i.e. Staré Sedlo).

1970a Trigonobalanus exacantha MAI , p. 385, pl. 1, figs 19-26, pl. 2, 1-15, pl. 3, figs 3-6, 15-19 (Wiesa).

1988 Trigonobalanopsis rhamnoides (ROSSMÄSSLER) KVAČEK et WALTHER , p. 405, pl. 2, figs 1-8, pls 50-54, pl. 55, figs 2-7, pl. 56, figs 1-4, pl. 57, text-figs 2-4 (Staré Sedlo, Wackersdorf Hrádek/N., Kristina Mine, Wiesa, Kleinsaubernitz, Plesná, drill core V 146).

1988 Trigonobalanopsis exacantha (MAI) KVAČEK et WALTHER , pp. 404, 413, pls 47-48, pl. 49, fig.8, pl. 55, fig.1 (Hrádek/N. Kristina Mine, Wiesa, Cheb Basin, drill core V 14).

1989 Trigonobalanopsis rhamnoides (ROSSMÄSSLER) KVAČEK et WALTHER ; Kvaček and Walther, p. 224, figs 4g-j, l (Jehličná, Wiesa, Hrádek/N., Kristina Mine, Adendorf).

1989 Trigonobalanopsis exacantha (MAI) KVAČEK et WALTHER ; Kvaček and Walther, p. 222, figs 4a-f, k (Hrádek/N., Kristina Mine, Wiesa).

Leaves petiolate, mostly incompletely and fragmentarily preserved, lamina elliptic to ovate, 35–50 mm long and 10–20 mm wide, base cuneate to rounded with short petiole (up to 7 mm long), apex attenuate and obtuse, margin entire, venation brochidodromous, midrib strong, straight, secondary veins thinner, distinct, straight or curved, looping at margin, alternate, regularly spaced, originating at an angle of 25–40°, tertiary veins alternate to opposite percurrent, straight to sinuous, rarely forked, venation of higher orders regular polygonal reticulate, areolation well developed, areoles 4-sided, veinlets one branched. Adaxial epidermis thinly cutinized, reflecting outline of polygonal cells 10–15 µm in size, anticlinal walls straight (in sun leaves) or minutely undulate (in shade leaves). Trichome bases sparsely distributed. Abaxial epidermis medium cutinized, showing non-modified polygonal cells 10–15 µm in size, rounded cyclocytic stomata 20–25 µm in diameter with 3–4 narrow and thickened subsidiary cells, variously dense simple bases of glandular serial trichomes and on the thick veins solitary broader thickly cutinized bases left by unicellular trichomes. Differences of shade and sun leaves are expressed in the intensity of sinuous anticlinal cell walls and density of stomata and glandular trichome bases (see Kvaček and Walther 1978). Cupules mostly with 3, rarely 2, 4 or 5 valves, narrow pyramidal to broadly ovoid, 9–15 mm long, 5–10 mm wide, at base cuneate, often with an oblique stalk, valves keeled, fused only in the lower third to half, outer surface coarsely striate, fruits pyramidal, trigonal, 4–7 mm long, 3–4.5 mm wide, widest in the lower third or half, often strongly compressed, edges sharp to slightly winged, sides convex, base rounded, apex with three widely open styles ( Holý 1975, p. 35, pl. 5, figs 11-14, as Trigonobalanus exacantha ,).

D i s c u s s i o n: Foliage of this extinct member of the Fagaceae has usually been recognized mainly according to the rhamnoid venation ( Phyllites rhamnoides ROSSM. ) and a very typical abaxial cuticle. First Kräusel and Weyland (1954) recognized its affinity to the Fagaceae and assigned it to Castanopsis . Ecotypical variation of sun and shade leaves led them to recognize two species, Castanopsis dechenii (WEBER) KRÄUSEL et WEYLAND and C. toscana (BANDULSKA) KRÄUSEL et WEYLAND. Kvaček (1966) suggested a similar interpretation for the material from the Kristina Mine. However a parallel comparative study of fruits and leaves from several occurrences in the European Tertiary led Kvaček and Walther (1988, 1989) to establish an independent fossil genus Trigonobalanopsis for this kind of foliage as well as for fruits originally interpreted as Trigonobalanus by Mai (1970a) and regularly associated together. The fruits assigned to Trigonobalanopsis exacantha (MAI) KVAČEK et WALTHER are very common in the Hrádek flora ( Holý 1977a, p. 112, as Trigonobalanus exacantha MAI, Kvaček and Walther 1988, p. 41 , pl. 60, figs 2-7, 1989, text-figs 4 a-b, Teodoridis 2003, pp. 14-15, pl. 3, figs 5-8).

The taxonomically nearest living relative is certainly Trigonobalanus doichangensis (A. CAMUS) FORMAN from SE Asia ( Laos, southern China, etc.), recently removed from this genus to Formanodendron NIXON et CREPET (1989) . However, Kvaček (2007) rather saw the nearest ecological analogue of Trigonobalanopsis in several living species of Castanopsis from southern Asia.

M a t e r i a l: Leaf compressions on slides, G 7698, 9195-9265 (KR 6, 37, 42, 54, 55, 72, 78, 81, 87, 88, 92, 96, 98, 99, 133, 134, 137, 138, 145, 154, 158, 165, 195, 196, 214, 301, 302, 304, 305, 306, 307, 308, 309, 310, 311, 312, 314, 315, 316, 383, 500), numerous cupules and fruits, G 3010, 4622, 5403, 8818-20, Gs 100, 102, 233-235.

Quercus L. Quercus bavarica (E. KNOBLOCH et KVAČEK) KVAČEK Pl. 3, fig. 5, pl. 13, fig. 1

1976

2004

Castanopsis bavarica E. KNOBLOCH et KVAČEK , p. 40, pl. 4, figs 2-3, 9-10, pl. 15, fig. 6, pl. 21, figs 1-4, text-fig. 15 (Wackersdorf).

Quercus bavarica (E. KNOBLOCH et KVAČEK) KVAČEK , p. 12, pl.10, figs 1-5, 12-13 (Flörsheim).

Leaf incomplete, elliptic, 23 mm long, 11 mm wide, base cuneate, apex not preserved, margin entire, venation eucamptodromous, midrib strong, straight, secondary veins thinner, distinct, curved towards the margin, alternate, relatively widely spaced, originating at an angle of 30–40°, tertiary veins poorly preserved, alternate percurrent, straight to sinuous or forked, venation of higher orders regular polygonal reticulate, areolation well developed, areoles 4-sided. Adaxial epidermis thinly cutinized, smooth to granulate outside, non-modified cells polygonal, ca. 12 µm in diameter, anticlinal walls straight, abaxial epidermis medium cutinized, smooth except for occasional granulate periphery of stomata, non-modified cells straight-walled, 10–12 µm in diameter, stomata cyclocytic, subsidiary cells 5–6, very little different from the non-modified cells, guard cell pairs elliptical, 20–22 µm long and 12–15 µm wide, with polar very thin T pieces, stomatal ledges broad, moderately thickened around small elliptical pore, trichome bases quite sparse, rounded, 15 µm in diameter, with subsidiary cells reaching under the base and leaving small polygonal scar of terminal tríchome part 8 µm in diameter, often asymmetrically positioned. Terminal parts of trichomes not preserved.

D i s c u s s i o n: Kvaček (2004) in agreement with others ( Kvaček and Walther 1988, Walther 1999, Meller et al. 1999) was in favour of transfering this rare element of the mastixioid floras from Castanopsis to Quercus L. More comparative studies are required to fix the systematic position of Quercus bavarica within this genus.

M a t e r i a l: Incomplete leaf compressions on slide, G 9266-9267 (KR 23, 496).

Quercus rhenana (KRÄUSEL et WEYLAND) E. KNOBLOCH et KVAČEK plexus

Pl. 3, figs 8-10, pl. 7, figs 3-4, pl. 13, figs 3-4

1950 Illicium rhenanum KRÄUSEL et WEYLAND , p. 50, pl. 9, figs 5-7, pl. 10, figs 1-2, pl. 11, fig. 5 (Brühl).

1966 Quercus lusatica JÄHNICHEN , p. 478, pls 1-4, pl. 9, figs 21-22, text-figs 1, 3-4 (Hartau, Wiesa).

1976 Quercus rhenana (KRÄUSEL et WEYLAND) E. KNOBLOCH et KVAČEK , p. 41, pl. 17, figs 6, 8, 14, pl. 21, figs 5-6, pl. 34, fig. 10 (Wackersdorf, Oder 2a Mine).

1999b Quercus sp. ? Sect. Phellos LOUD.; Mai, p. 34, pl. 11, fig. 3 (Wiesa).

1999b Quercus sapperi (MENZEL) MAI ; Mai, p. 33, pl. 11, fig. 6 (Hartau).

For more synonyms and discussion see Jähnichen (1966, as Quercus lusatica ).

Incomplete leaves and fragments, lamina elliptic to ovate, 14–78 mm long and 7–29 mm wide, apex shortly attenuate and blunt, base cuneate to widely cuneate, margin entire, venation brochidodromous to eucamptodromous, midrib strong, straight, secondary veins thinner, regularly spaced, alternate to subopposite, straight, curved towards the margin, originating at an angle of 40–60°, tertiary veins alternate percurrent, straight to sinuous or often forked, venation of higher orders regular polygonal reticulate, areolation well developed, areoles 4-sided, veinlets lacking. Adaxial epidermis medium thick, cuticle fine punctate on surface showing polygonal non-modified cells with seemingly straight, but in detail fine undulate anticlines, rare solitary simple trichome bases with a ring of small subsidiary cells, abaxial epidermis medium cutinized, showing polygonal outlines of non-modified cells and large rounded cyclocytic stomata with a pair of guard cells 28–30 µm in diameter surrounded by a ring of massively cutinized subsidiary cells with strong striation, which may partly be shared by two adjacent stomata. Simple rounded trichome bases ca. 15 µm in diameter. Acorns ovoid, partly compresed, 23–43 mm long and 17–29 mm wide, without cupule, base rounded, apex shortly attenuated and blunt, apical surface with fine paralel striation.

D i s c u s s i o n: Foliage of this oak commonly occurs in coaly facies in several sites in the Miocene of Europe. Because the gross morphology is not very characteristic and can be easily mistaken for Lauraceae , it is difficult to decide on the correct name (see e.g., Palamarev and Mai 1998, p. 250-251, Quercus lyellii group). Even the epidermal characteristics vary among similar leaf forms and may be useful in dividing various fossil populations over Europe (cf. Kovar-Eder 1996). The occurrence from the Kristina Mine matches the records from the neighbouring sites of Turów in Poland (as Magnolia sp. sensu Czeczott et al. 1959) and Saxony (as Castanopsis sp. sensu Mai 1964). The leaves are regularly accompanied at Hartau, Turów ( Jähnichen 1966, p. 507), Hrádek/N., and Medard Mine by fruits and cupules more recently assigned to Quercus sp. (? Sect. Phellos LOUD.) by Mai (1999b, p. 34, pl. 11, fig. 3) and the Quercus sapperi (MENZEL) MAI type ( Mai 1999 b, p. 33, pl. 11, fig. 6).

Because of quite unusual epidermal pattern, Kvaček (1966) hesitated to use a modern genus and called the foliage of this element Quercophyllum sp. Indeed we could speculate that Quercus rhenana may represent an extinct entity within the Fagaceae subfam. Quercoideae but the complete characteristics, including in particular male inflorescences and pollen in situ are needed to resolve its relationship.

M a t e r i a l: Fragmentary leaf compressions on slides, G 9268-9285 (KR 57, 75, 93, 94, 95, 152, 156, 167, 184, 215, 467-473), compression of fruit, G 9286 (KR 175), impressions of leaves, G 8814a, b, and a fruit, G 8813, in ironstone.

Juglandaceae DC ex PERLEB.

Engelhardia LESCHEN. ex BLUME

Engelhardia orsbergensis (WESSEL et WEBER) JÄHNICHEN, MAI et WALTHER

Pl. 3, figs 10-14, pl. 13, fig. 5

1856

1977

Banksia orsbergensis WESSEL et WEBER , p. 146, pl. 25, figs 9a-d (Orsberg).

Engelhardia orsbergensis (WESSEL et WEBER) JÄHNICHEN, MAI et WALTHER , pp. 326-346, pls 38-49, text-figs 1-3 (localities and stratigraphy see Jähnichen et al. 1977, pp. 336-337).

Complete leaves not preserved, leaflets sessile, shortly petiolulate, lamina elongate to narrow elliptic, 20–60 mm long, 10–20 mm wide, apex acute, base asymmetric, rounded to widely cuneate, margin simple coarsely to minutely serrate, teeth acute, sinus angular, in the lower leaflet part entire, venation semicraspedodromous, midrib strong, straight to slightly curved, secondary veins numerous, distinctly thinner, mostly straight, alternate, originating at an angle of 50–80°, looping at the margin, intersecodaries parallel, thinner, tertiary veins percurrent, straight to sinuous, venation of higher orders regular polygonal reticulate, areolation well developed, 3- to 4-sided, veinlets lacking. Abaxial epidermis thinly cutinized, composed of non-modified polygonal rounded cells, poorly preserved, stomata anomocytic, widely elliptic, 20–26 µm long and 15–18 µm wide, sunken and usually seen only as traces of outer ledges surrounded by a circle of low papillae, peltate trichomes rarely preserved with large lens-shaped head 130–150 µm and more in diameter, crenulate on margin, composed of numerous narrow segments, trichome bases of uniform size, rounded, typically with double outline, 19–24 µm in diameter.

D i s c u s s i o n: The leaf records of this typical extinct representative of the Juglandaceae , tribe Engelhardiae ( Manchester 1987) have been variously assigned to Engelhardia ( Saporta 1865, Kvaček 1972 as Engelhardia detecta SAPORTA, Jähnichen et al. 1977 as Engelhardia orsbergensis ), Palaeocarya SAPORTA ( Jähnichen et al. 1984, Manchester 1987) and Oreoroa DILCHER et MANCHESTER (1986) . The obtained leaflets, although incomplete, match in all respects the other occurrences of this sort in Europe, e.g., from Wackersdorf ( Knobloch and Kvaček 1976) and elsewhere ( Jähnichen et al. 1977). The currently employed epithet is, however, illegitimate (a later synonym – see Winterscheid and Kvaček, personal communiatioín) and requires conservation.

M a t e r i a l: Leaflet compressions on slides, G 9287- 9306 (KR 97, 168, 202, 212, 213, 217, 218, 219, 253, 479, 525).

Pterocarya KUNTH. Pterocarya limburgensis C. et E.M. REID Pl. 7, figs 6-7

1915 Pterocarya limburgensis C. et E.M. REID, p. 73, pl. 4, figs 15-21 (Reuver).

1961 Pterocarya cf. stenoptera DC ; Czeczott and Skirgiełło, pp. 54, 104, pl. 16, figs 1-6 (Turów).

1964 Pterocarya cf. raciborskii ZABLOCKI ; Mai, p. 103-104, pl. 13, fig. 7 (Hartau).

1977a Pterocarya raciborskii ZABLOCKI ; Holý, p. 112 (Hrádek/N., Kristina Mine).

2003 Pterocarya limburgensis . C. et E.M. REID; Teodoridis, p. 16, pl. 3, figs 11-12 (Hrádek/N., drill cores).

Nuts obliquely ovoid to broadly ovoid, transversally elongate, variable in form, strongly compressed, 3.2–5.3 mm tall, 3.0–5.3 mm wide, base obliquely bent, usually rounded, apex shortly pyramidal, pointed, rarely long elongate, bent, with remnants of style, 10 (9–12) strong wing ribs often anastomosing or forked running from the base, more distinct at the base and the lower third or half, disappearing towards but never entering the apex, dehiscence line on a meridional rib running over the apex, inner structures not known ( Holý 1975, pp. 31-32, pl. 5, figs 8-10).

D i s c u s s i o n: Holý (1975, 1977a) maintained the morphological identity of the material from the Kristina Mine with the poorly illustrated type material of Pterocarya raciborskii ZABLOCKI from Wieliczka in Poland ( Zablocki 1928, p. 189, pl. 10, figs 21-34 – see Łancucka-Środoniowa and Zastawniak 1997) based only on the similar size variation. Nevertheless the above-described material corresponds to Pterocarya remains indicated from the Zittau Basin and Saxony ( Czeczott and Skirgiełło 1961, Mai 1964, 1999b, Czaja 2003, Teodoridis 2003). The nearest living relative appears to be Pterocarya hupehensis SKAN. Mai (1999b, p. 46) includes Pterocarya raciborskii into the genus Cyclocarya ILJINSKAYA.

M a t e r i a l: More than 50 fruits, G 3065, 8977-78, Gs 84.

Cyclocarya ILJINSKAYA Cyclocarya nemejcii HOLÝ

Pl. 7, figs 8-9

1977a Cyclocarya nemejcii HOLÝ , p. 114, pl. 2, figs 1-14 (Hrádek/N., Kristina Mine).

For the description and discussion see Holý (1977a, p. 114). Mai (1999b, p. 46) stresses a great similarity with Cyclocarya nucifera (LUDWIG) MAI and suggests undertaking statistical morphometrical studies for discrimination of the two species.

M a t e r i a l: More than 20 endocarps, G 3049, 4307-09.

Myricaceae A. RICH. ex KUNTH

Myrica L. Myrica ceriferiformis KOWNAS

Pl. 7, figs 10-11

1955 2003 Myrica ceriferiformis KOWNAS , p. 459, figs 8 a-b (Dobrzyn). Myrica ceriferiformis KOWNAS ; Teodoridis, p. 16, pl. 3, figs 13-21, pl. 4, figs 1, 4 (Hrádek/N., drill cores).

Endocarps secondarily compressed, medium thin-walled, roundish to ovoid, 3–4.3 mm long, 2.6–3.7 mm wide, dehiscence sutures wide, without leaving a rib on the endocarp surface, apex with a tiny point (remain of the style), exocarp composed of ca. 12–14 tightly packed tiny verrucae ( Holý 1975, p. 25, pl. 4, figs-1-2).

D i s c u s s i o n: According to Holý (1975) the material matches the type specimens from the Polish locality Dobrzyn ( Kownas 1955) except for finer verrucae. Fruits from Klettwitz and Piskowitz ( Mai 1964) are usually ovoid, with fewer verrucae on the surface than is seen in the type illustrated in Kownas (1955). Similar fruits are produced by Myrica cerifera L. and M. pensylvanica LOIS. living in the Atlantic North America. Similar endocarps and exocarps have been decribed from the drill cores near Hrádek/N. as Myrica ceriferiformoides BŮŽEK et HOLÝ ( Teodoridis 2003) . In the case that the exocarp is preserved, both taxa are clearly distinguished, however the endocarps of M. ceriferiformis , M. ceriferiformoides as well as M. suppanii KIRCHH. and M. boveyana (HEER) CHANDLER show very similar morfological charactersitics allowing mistakes to be made in their specific determinations (e.g., Bůžek and Holý 1966, Gregor 1975b, Teodoridis 2003). We are sure that the same also applies to endocarps of the two next species of Myrica defined by Holý (1977a, 1978a).

M a t e r i a l: 10 endocarps, G 8861-62.