Tectocarya robusta KIRCHHEIMER

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 : 80-83

cf. Tectocarya robusta KIRCHHEIMER

Pl. 9, fig. 1

?1935b Tectocarya robusta KIRCHHEIMER , p. 67, pl. 8, fig. 24a, f (Merka).

1977b cf. Tectocarya robusta KIRCHHEIMER ; Holý, p.138, pl. 4, figs 20-28 (Hrádek/N., Kristina Mine).

For the description and discussion see Holý (1977b). M a t e r i a l: 10 endocarps, G 4186-91.

Retinomastixia KIRCHHEIMER Retinomastixia oertelii GREGOR Pl. 8, figs 14-15

1977b Retinomastixia schultei KIRCHHEIMER ; Holý, p. 134, pl. 4, figs 27-32 (Hrádek/N., Kristina Mine).

1978b Retinomastixia oertelii GREGOR , p. 64, pl. 14, figs 1-4 (Wackersdorf).

1999c Retinomastixia oertelii GREGOR ; Mai, p. 61, pl. 16, figs 14-16 (Sandförstgen, Hartau).

For further synonyms see Gregor (1978b)

For the description see Holý (1977b, as Retinomastixia schultei ), for the discussion Gregor (1975, 1978a) and Mai (1999c).

M a t e r i a l: Ca. 30 fruits, numerous fragments, G 3009, 4257-62.

Ericaceae JUSS.

Leucothoë D. DON Leucothoë narbonnensis (SAPORTA) WEYLAND plexus

Pl. 9, figs 2-3

1865 Andromeda (Leucothoë) narbonnensis SAPORTA , p.142, pl. 8, fig.1 (Armissan).

1943 Leucothoë narbonnensis (SAPORTA) WEYLAND , p. 118 (Armissan) (? non pl. 21, figs 3-6 – Rott).

1960 Leucothoë narbonnensis (SAPORTA) MAI , p.85, pl. 6, figs 5-16 (Wiesa).

1977a Leucothoë sp. ; Holý, p. 112 (Hrádek/N., Kristina Mine).

Pentamerous capsules 2.1–3.5 mm long, 1.4–2.9 mm wide, ellipsoid pear-shaped, short stalked at base, apex 5- sided, with a pit in the middle, fruits septicidal, smooth, dehiscence starting at the apex separating individual endocarps from the central stylar column, seeds bent elongate ovoid, 0.7–1.2 mm long, 0.5–0.8 mm wide, flattened on sides, with distinct meshes of prosenchymatous tissue of testa on surface ( Holý 1975, p. 51, pl. 8, figs 6-11).

D i s c u s s i o n: Holý (1975, p. 52) intended to separate fruits usually treated as Leucothoë narbonnensis under an independent fossil species, because Weyland (1943) allegedly pre-occupied the epithet “narbonnensis ” for vegetative remains. We disagree with this view (see also Mai 2000) and treat Leucothoë narbonnensis in sense of combined fossil species as “plexus”.

M a t e r i a l: Numerous fruits, G 3061, 8853-54, Gs 81.

Pentaphylacaceae ENGL. (incl. Ternstroemiaceae MIRB. ex DC )

Ternstroemia NUTIS ex L. Ternstroemia chandlerae HOLÝ

Pl. 9, figs 4-5

1977a Ternstroemia chandlerae HOLÝ ( “ chandleri ”), p. 117, pl. 3, figs 8-13 (Hrádek/N., Kristina Mine).

For the description and discussion see Holý (1977a). According to Mai (1999b, p. 35), this species is most probably synonymous with variable Ternstroemia sequoioides (ENGELHARDT) BŮŽEK et HOLÝ in BŮŽEK et al. (1996).

M a t e r i a l: Seeds, G 4300-04, 4612.

Eurya THUNB. Eurya stigmosa (LUDWIG) MAI Pl. 9, fig. 6

1860

1957 1960

1960

1961

Potamogeton stigmosus LUDWIG , p. 60, pl. 8, fig. 13 (Salzhausen).

Myrtospermum variabile CHANDLER , p. 112 (Bovey Tracey). Eurya stigmosa (LUDWIG) MAI , p. 79, pl. 4, figs 8-17 (Wiesa).

? Cleyera? variabilis (CHANDLER) CHANDLER , pp. 213, 225, pl. 31, figs 48-56, pl. 34, fig. 145 (Bournemouth).

Myrtospermum warrenii CHANDLER , p. 81, pl. 8, figs 10-19 (Reading beds).

1971 Eurya stigmosa (LUDWIG) MAI ; Mai, p. 329, pl. 34, figs 27-28 (Salzhausen and further references).

1977a Eurya stigmosa (LUDWIG) MAI ; Holý, p. 112 (Hrádek/N., Kristina Mine).

2003 Eurya stigmosa (LUDWIG) MAI ; Teodoridis, p. 19, pl. 4, figs 5, 7, 8 (Hrádek/N., drill cores).

Seeds thick, discoid, horseshoe-shaped, roundish reniform, variously obliquely asymmetric, very variable in shape, 1.3–2.0 mm in size, sides variously vaulted, flattened or concave, surface ornamented by ca. 10 concentric rows of radially elongate cells, 0.05–0.06 mm wide in honeycomblike structure, base straight or slightly cordate, shorter than half of the seed length, dehiscence in the plane of symmetry, condylus elliptic, high above the seed centre, seen as transversal, differently structured elevation, funicular canal as triangular cavity between embryonal pit and hilum, testa double layered, outer thicker layer composed of horseshoe-shaped radially disposed sclereids, inner layer composed of small isodiametric sclereids.

D i s c u s s i o n: According to Holý (1975, pl. 8, figs 1-4) Eurya stigmosa is certainly an aggregate species because of the wide variation in seeds. Eurya japonica THUNB. used to be indicated as the nearest living relative.

M a t e r i a l: Ca. 50 seeds, G 3059, 4611, 4613, Gs 98.

Styracaceae DC et SPRENG.

Rehderodendron HU Rehderodendron custodum HOLÝ

Pl. 9, figs 7-8

1977a Rehderodendron custodum HOLÝ , p. 118, pl. 4, figs 1-20 (Hrádek/N., Kristina Mine).

For the description and discussion see Holý (1977a). The species has been accepted, although transitions to R. ehrenbergii (KIRCHHEIMER) MAI with larger fruits exist (see also Mai 2000, p. 47).

M a t e r i a l: Ca. 40 endocarps, G 4228, 4326-4344, Gs 80; and G 3006, 8835 identified as R. ehrenbergii .

Symplocaceae DESF. Symplocos JACQUIN

European Tertiary representatives were revised by Mai and Martinetto (2006) and their treatment is followed here, although Holý (1975, pp. 52-59, 1977a, p. 112) originally recognized more species.

Symplocos schereri KIRCHHEIMER

Pl. 9, fig. 9

1935a Symplocos schereri KIRCHHEIMER , p. 291, fig. 8 (Konzendorf).

1940 Symplocos wiesaensis KIRCHHEIMER , p. 288, fig. 5 (Wiesa).

1977a Symplocos schereri KIRCHHEIMER ; Holý, p. 112 (Hrádek/N., Kristina Mine).

2006 Symplocos schereri KIRCHHEIMER ; Mai and Martinetto, p. 21, pl.2, fig. 9, pl. 8, figs 2-7 (review of sites).

Endocarps 5–10 mm long, 3.5–6.5 wide, elongate ovoid, 2–3 locular, apex blunt, base narrowed, usually with three points, apical pit deep, wider than the half of the endocarp width, distinct longitudinal winged ribs irregularly wavy and irregularly thick, often radially dichotomizing on the surface ( Holý 1975, pp. 52-53, pl. 9, figs 1-8, partly identified as Symplocos wiesaensis ).

D i s c u s s i o n: In the Hrádek flora Holý (1975) recognized in addition to S. schereri also S. wiesaensis , admitting that the two morphotypes were connected with transitional forms. His view was confirmed by the treatment by Mai and Martinetto (2006) in merging the two and suggesting S. tanakae MATSUMURA from Japan as the nearest living relative.

M a t e r i a l: More than 20 endocarps, G 4582, 8981, Gs 37.

Symplocos casparyi LUDWIG

Pl. 9, figs 10-11

1857 Symplocos casparyi LUDWIG , p. 99, pl. 20, figs 6a-f (Dorheim).

1860 Carpinus salzhausensis LUDWIG , p. 100, pl. 33, fig. 8 (Salzhausen).

1867 Carpolithus lignitarum QUENSTEDT , p. 914, pl. 86, fig. 35 (Salzhausen).

1935c Symplocos salzhausensis (LUDWIG) KIRCHHEIMER , pp. 718, 737, fig. 19 (Salzhausen).

1935c Symplocos lignitarum (QUENSTEDT) KIRCHHEIMER , p. 718, figs 17-18 (Salzhausen).

1977a Symplocos lignitarum (QUENSTEDT) KIRCHHEIMER ; Holý, p. 112 (Hrádek/N., Kristina Mine).

1977a Symplocos salzhausensis (LUDWIG) KIRCHHEIMER ; Holý, p. 112 (Hrádek/N., Kristina Mine).

2003 Symplocos lignitarum (QUENSTEDT) KIRCHHEIMER ; Teodoridis, p. 21, pl. 5, figs 8-10 (Hrádek/N., drill cores).

2003 Symplocos salzhausensis (LUDWIG) KIRCHHEIMER ; Teodoridis, pp. 21-22, pl. 5, figs 11, 13, 14 (Hrádek/N., drill cores).

2003 Symplocos lusatica MAI ; Teodoridis, p. 21, pl. 5, fig. 12 (Hrádek/N., drill cores).

For further synonyms see Mai and Martinetto (2006).

Endocarps variable in form, two forms are recognizable in the Kristina flora: “ lignitarum ” – endocarps 4.1– 9 mm long, 3.3–4.8 mm wide, mostly short cylindrical to elongate ovoid, rarely bent, mostly trilocular (exceptionally with 2 or 4 locules) with convex sides, base rounded, often with a small central pit, apex straight, truncate, apical pit shallow reaching half of the width of endocarps, surface with longitudinal shallow blunt ribs running throughout the length of the endocarps, rarely smooth, “ salzhausensis ” – endocarps 2.5– 5 mm high, 3.6–4.8 mm wide, widely ovoid to transversally ovoid, base rounded or blunt, often with a central pit, apex rounded, blunt, apical pit shallow, rounded to triangular, reaching to one third of the complete width of endocarps, mostly trilocular with evenly developed locules, on surface often smooth or with indistinct low rounded widely spaced ribs or elongate protuberances ( Holý 1975, pp. 54-57, pl. 9, figs 9-12, pl. 10, figs 1-7, as Symplocos lignitarum and S. salzhausensis ).

D i s c u s s i o n: According to Holý (1975) the “ lignitarum ” morphotype is most common in central Europe (Hartau, Turów) and also in the Hrádek flora. Mai and Martinetto (2006) recognized still more morphotypes and indicated other living species (e.g., S. glandulifera BRAND) matching in fruit morphology.

M a t e r i a l: Numerous endocarps (G 4583, 4631, 6493, 8982-83, Gs 85, Gs 86).

Symplocos pseudogregaria KIRCHHEIMER

Pl. 9, fig. 12

1938a Symplocos pseudogregaria KIRCHHEIMER , p. 354, pl. 7, figs 17-20, pl. 8, figs 1-2 (Niederpleis).

1941a Symplocos poppeana KIRCHHEIMER , p. 217, figs 12-13 (Wiesa).

1977a Symplocos poppeana KIRCHHEIMER ; Holý, p. 11 (Hrádek/N., Kristina Mine).

Endocarps 5.4– 9 mm long, 3–4.1 mm wide, trilocular, mostly cylindrical, base rounded, apex truncate, with a wide, shallow apical pit, surface cover by fine longitudinal striae and rows of protuberances or furrows; 3 of them following the sutures of carpels from the base to about half of the endocarp length ( Holý 1975, pp. 57-58, pl.10, fig. 8, as Symplocos poppeana ).

D i s c u s s i o n: Mai and Martinetto (2006) reinterpreted Symplocos pseudogregaria as a polymorphic species and merged with it several morphotypes, including Symplocos poppeana , which was indicated for the Hrádek flora by Holý (1975, 1977a). They listed several living species having similar endocarps, e.g., S. anomala BRAND, S. tingifera CHEN and S. kuroki NAGAMASU , distributed in E Asia (southern Japan, Yunnan, Indonesia).

M a t e r i a l: 5 poorly preserved endocarps, G 8984, Gs 89.

Symplocos minutula (STERNBERG) KIRCHHEIMER Pl. 9, fig. 13

1825 Carpolithes minutulus STERNBERG , p. 41, pl. 53, fig. 8 (“ Carpolites ”) (Salzhausen).

1949 Symplocos minutula (STERNBERG) KIRCHHEIMER , p. 16, pl. 1, fig. 6, pl. 2, fig. 16 (Salzhausen).

1977a Symplocos minutula (STERNBERG) KIRCHHEIMER ; Holý, p. 112 (Hrádek/N., Kristina Mine).

Endocarps 5.5–7.7 mm long, 3.0–4.7 mm wide, slightly bent, elongate ovoid, (?) trilocular, base rounded, sides towards apex narrowed, apical pit deep, enclosed by the margin, on the neck often corroded and split, surface smooth, at most very thinly striate-wrinkled ( Holý 1975, pp. 68-69, pl. 11, figs 1-3).

D i s c u s s i o n: The Sternberg’s collection at NM includes syntypes of Carpolithes minutulus ( Holý 1975, Kvaček and Straková 1997), contrary to the statement of Kirchheimer (1957). However, the lectotypification has not been done. Mai and Martinetto (2006) listed S. tinctoria L´ HÉRITIER (SE USA) and S. ramossissima WALL. (Himalayas) and S. glauca (THUNBERG) KOIDZUMI ( Japan) as living species with similar endocarps.

M a t e r i a l: 6 compressed endocarps, G 8985, Gs 88.

Sphenotheca KIRCHHEIMER Sphenotheca incurva KIRCHHEIMER

Pl. 9, fig. 14

1934 Sphenotheca incurva KIRCHHEIMER , p. 789, fig. 19 (Kausche). 1977a Sphenotheca incurva KIRCHHEIMER ; Holý, p. 112 (Hrádek/N.,

Kristina Mine).

Fruits sack-shaped, elongate, slightly asymmetrical 14.3–15.5 mm long, 7.2– 9 mm in diameter, neck-like narrowed below slightly oblique truncate apex with a broad collar and wide central pit with canal openings from three locules, base rounded, on one specimen mesocarp abraded to the surface of longitudinal pectinate wings of endocarp ( Holý 1975, p. 59, pl. 11, fig. 4).

D i s c u s s i o n: This extinct member of the Symplocaceae is a marker of the Younger Mastixioid Floras of Europe ( Mai 2000).

M a t e r i a l: 2 fruits, G 4584, 5409.

Theaceae MIRB. ex KER GAWL.

Gordonia ELLIS

Gordonia hradekensis (KVAČEK et BŮŽEK) BOZUKOV et PALAMAREV

Pl. 3, figs 26-30, pl. 14, figs 4-5

1966 Symplociphyllum hradekense KVAČEK et BŮŽEK , p. 293, pl. 2, figs 5-6, pl. 3, fig. 1, pl. 4, figs 8-9 (Hrádek/N., Kristina Mine).

1984b Polyspora hradekensis (KVAČEK et BŮŽEK) KVAČEK et WALTHER , p. 335, pls 57-59 (Hrádek/N., Kristina Mine, Wackersdorf).

1995 Gordonia hradekensis (KVAČEK et BŮŽEK) BOZUKOV et PALAMAREV , p. 182, text-fig. 7 (Satovcha).

Leaves simple, lamina oblong to elliptic or slightly ovate, up to 33–62 mm long and 9–14 mm wide, base cuneate with petiole, up to 10 mm long, apex attenuate to acute, rarely emarginate, margin entire in basal part, upper margin regularly simply serrate, teeth blunt to acute, glandular, sinus angular to rounded, venation semicraspedodromous, midrib strong, straight, secondary veins thinner, curved, alternate, rarely opposite, originating at an angle of 30–50°, tertiary veins alternate to opposite percurrent, straight to sinuous or forked, venation of higher orders regular polygonal reticulate, areolation well developed, areoles 3- to 4-sided, veinlets unbranched. Adaxial epidermis thickly cutinized, smooth, reflecting outlines of non-modified cells 30 µm or more in diameter, anticlinal walls curved to undulate with lens-shaped thickenings in sinuses, solitary star-like simple trichome bases 20–25 µm in diameter and with thickened outer wall, abaxial epidermis medium cutinized, strongly striate around the stomata, non-modified cells similar to those of the adaxial epidermis, stomata anisocytic to cyclocytic, guard cell pairs rounded to broadly oval, 25–35 µm wide and 30–40 µm long, trichome bases of the same type as in the adaxial epidermis, scattered over the whole surface.

D i s c u s s i o n: Kvaček and Walther (1984b) corrected the previous view of Kvaček (1966, in Kvaček and Bůžek 1966) who erroneously interpreted the stomatal type of Symplociphyllum hradekense KVAČEK et BŮŽEK as paracytic suggesting affinities with Symplocaceae , and recognized Polyspora SWEET as the nearest living relative based on wide comparative study ( Kvaček and Walther 1984a). This genus is generally, but not always ( Judd et al. 2002) considered a later synonym of Gordonia ELLIS. Bozukov and Palamarev (1995) supplied the resulting combination. A transfer to Styrax ( Styracaceae ) ( Schweigert 1992) is not supported by the epidermal anatomy and hence not acceptable. The fossil genus Symplociphyllum KVAČEK et BŮŽEK must be re-typified, because its original type falls into the genus Polyspora or Gordonia (see below).

M a t e r i a l: Several mostly incomplete leaf compressions on slides, G 9328-9389 (KR 28, 29, 30, 35, 38, 44, 47, 48, 49, 64, 69, 70, 71, 86, 96, 153, 160, 170, 172, 173, 185, 192, 207, 210, 211, 220, 222, 223, 229, 232, 239, 249, 250, 251, 252, 299, 429, 435, 524).

Oleaceae HOFFMANS et LINK

Fraxinus L. Fraxinus bilinica (ETTINGSHAUSEN) KVAČEK et HURNÍK

Pl. 4, figs 1-4, pl. 14, figs 6-7

1849

1860

2000

Juglans bilinica UNGER , p. 126 pro parte (non pl. 14, fig. 20) (Bílina).

Carya bilinica (UNGER) ETTINGSHAUSEN ; Unger, p. 39, pro parte, pl. 17, figs 1-8 (Bílina).

Fraxinus bilinica (UNGER) KVAČEK et HURNÍK , p. 19, pl. 8, figs 7(-?8), text-figs 4.2, 4.7 (Zabrušany, Louny-Vršovice).

Leaflets elliptic, 12–33 mm long, 9–35 mm wide, apex not preserved, probably short acuminate, base cuneate, margin bluntly serrate, teeth small, blunt, sinus angular, venation semicraspedodromous, midrib strong, straight, secondary veins thinner, almost straight to slightly curved, alternate, rarely subopposite, originating at an angle of 40–60°, looping near margin, occasional intersecondaries thin, short, parallel and sinuous, tertiary veins alternate percurrent, straight to sinuous, often forked, innervating mostly teeth sinuses on margin, venation of higher orders regular polygonal reticulate, areolation well developed, areoles 3- to 4-sided, veinlets lacking. Leaf lamina tough, but epidermis thinly cutinized thus the cuticle structure hardly observable, clearest in sub-macerated compressions. Adaxial cuticle indistinctly striate to punctate, reflecting outlines of polygonal cells 20–25 µm in diameter with straight or very little wavy anticlinal walls, on veins slightly elongate, with rounded simple bases 6–8 µm in diameter formed after small peltate glandular trichomes. Abaxial cuticle still thinner, smooth, showing only scattered pairs of guard cells 20–30 µm long and 10–20 µm wide with slightly thickened boat-like stomatal ledges and trichome bases of the same type as adaxially, but denser, occasionally with a rarely preserved rounded polycellular shield 25–50 µm in diameter.

D i s c u s s i o n: Kvaček (1966) separated the Hrádek ash foliage with almost straight adaxial epidermal anticlines (as Fraxinus sp. ) from a younger population from the Pannonian of South Moravia at Poštorná, which differs in its much more undulate adaxial anticlines. This difference was also confirmed by Kovar-Eder (in Kovar-Eder and Krainer 1991) who assigned the material of Poštorná to F. angusta HUMMEL. Kvaček and Hurník (2000) suggested uniting the Early Miocene occurrences of ash foliage from Bílina and elsewhere under the binomen Fraxinus bilinica (UNGER) KVAČEK et HURNÍK leaving aside the previously used name of F. ungeri (GAUDIN) E. KNOBLOCH et KVAČEK for priority reasons. According to Kovar-Eder (1996) the ash foliage from Hrádek identified here as F. bilnica clearly corresponds in leaf epidermal anatomy with the record from the Oberdorf Mine at Köflach. No living analogous species has been suggested so far.

M a t e r i a l: Isolated incomplete leaflet compressions on slides, G 9381-9397, partly missing (KR 39, 40, 41, 60, 80, 91, 191, 411, 414, 415).

Araliaceae JUSS. Araliaceae gen. et sp. indet.

Pl. 9, figs 15-16

Endocarps (?) quite flat, almost half-moon to asymmetrically elongate to broadly ovoid, base and apex broadly rounded, the latter partly narrowed, ventral side straight to slightly convex, dorsal side almost semicircular, surface of lateral sides smooth, lustrous, sometimes with bent transversal striation ( Holý 1975, p. 93, pl. 80, figs 8-9).

D i s c u s s i o n: According to Holý (1975), similar but smaller endocarps are produced by Aralia L., while in Schefflera J.R. FORST. et G. FORST. or Eleutherococcus MAXIM. (syn. Acanthopanax (DECNE. etPLANCH.) MIQ.) they are similar in size, but with different sculptures. The lack of comparative living material prevented even the generic identification. The specimens are not available for further studies.

M a t e r i a l: 8 endocarps, missing.

Aquifoliaceae DC ex A. RICH.

Ilex L. Ilex cf. protogaea MAI

Pl. 9, fig. 17

?1970b Ilex protogaea MAI , p. 459, pl. 62, figs 9-15 (Schlabendorf).

Stone slender, 4 mm long, 1.2 mm wide, in lateral view long boat-like, ventral edge straight, sharp, sides joining under a sharp angle, bearing two very thin longitudinal ribs, dorsal side narrow, with two longitudinal anastomosing ribs ( Holý 1975, p. 66, pl. 12, figs 13-14).

D i s c u s s i o n: According to Holý (1975), the relationship to the similar Ilex cantalensis E.M. REID and I. lusatica MENZEL are discussed in Mai (1970b, p. 459). A living species with similar endocarps appears to be I. dahoon WALT.

M a t e r i a l: One stone, G 3062.