The Conifer Brachyphyllum Squammosum from the Bohemian Cenomanian

Acta Palaeobotanica 47(1): 25–35, 2007

The conifer Brachyphyllum squammosum from the Bohemian Cenomanian

JIŘÍ KVAČEK

National Museum, Prague, Václavské nám. 68, 115 79 Praha 1, Czech Republic; e-mail: [email protected]

Received 21 May 2006; accepted for publication 15 November 2006

ABSTRACT. Brachyphyllum squammosum (Velenovský) Palibin from the Bohemian Cenomanian (Czech Repub- lic) is revised. New material, represented by conifer twigs, from the locality of Pecínov (60 km west of Prague) is described including the well-preserved micromorphology of its epidermis. Similar Brachyphyllum species from the Cretaceous and Jurassic are discussed. A marked similarity was recorded between B. squammosum and B. vulgare (Stopes & Fujii) Jeffrey from the Upper Cretaceous of Japan. The misinterpretation by Velenovský and Bayer of reproductive structure putatively associated with B. squammosum is brieﬂ y discussed. The lectotype of B. squammosum is designated. The palaeoecological conditions of growth for B. squammosum are brieﬂ y discussed.

KEY WORDS: Brachyphyllum, conifer, Cenomanian, Upper Cretaceous, Czech Republic

INTRODUCTION

The conifer Brachyphyllum squammosum (Fig. 1), is infi lled by Upper Cretaceous fresh- is a rather rare fossil plant of the Bohemian water, brackish and marine sediments of Cenomanian. It was described by Velenovský Cenomanian to Campanian age. (1885) from the locality of Vyšehořovice. For The Peruc-Korycany Formation is situated a long time this locality was its unique occur- in the basal most position of the Bohemian rence in the Bohemian Cenomanian. In the Cretaceous Basin. The locality of Vyšehořovice, seventies of the last century, Němejc and 30 km east of Prague, comprising several Kvaček (1975) recorded the species from the sandstone quarries that were a source of pal- Santonian of South Bohemia. Finally, in the aeobotanical material for nearly one century, period between 1993 and 2003, several com- is now completely abandoned and preserved pressed specimens of Brachyphyllum squam- as a Natural monument. The Pecínov quarry, mosum were recovered in the Pecínov quarry. situated 60 km west of Prague, is a work- Although its systematic affi nity remains ing quarry where the entire Peruc-Korycany unresolved, the material is interesting for its Formation is exposed. The sedimentary suc- well-preserved cuticle and overall rarity. Its cession in Pecínov was divided by Uličný and description and typifi cation is the topic of the Špičáková (1996) into 5 units. Units 1–2 typi- following report. cally include fl uvial pebbly sandstones, con- glomerates and sandstones with interbedded mudstones. Unit 3 consists of mudstones rich LOCATION, GEOLOGY in pyrite concretions. They are the products of AND MATERIAL marginal marine and brackish sedimentation in back swamps and in supratidal marshes. The Bohemian Cretaceous Basin, as defi ned Unit 4 is represented by cross-bedded sand- by Čech et al. (1980), located in the Bohemian stones, mudstones and laminites, products of Massif, the Czech Republic, Central Europe sedimentation on a tidal fl at crossed by mean- 26

Fig. 1. A – Location of the Bohemian Cretaceous Basin in Central Europe; dark grey area indicates the Cretaceous basin, light grey area indicates the Bohemian Massif. B – Location of the two fossil sites mentioned in this paper, Czech Republic (after Uličný et al. 1997) dering tidal creeks. The lower part of unit 5 is removed in a drop of distilled water on an emulsion sur- built of sandstones containing a rich marine face of pieces of glossy negative fi lm. The small sheets fauna and occasionally preserved stems of tree were air-dried and mounted on SEM stubs. Cuticle preparations were studied by light micro- ferns and poorly preserved leaf impressions. scopy using Nomarski DIC (Olympus BX50) and by Intertidal to supratidal mudstones bearing SEM (Tesla BS340). All studied material is housed in a rich megafl ora are locally preserved in the the collections of the National Museum, Prague. uppermost part of unit 5 (Uličný et al. 1997), and refl ect local regression. Detailed bios- tratigraphical studies based on pollen spectra SYSTEMATICS (Pacltová 1977) date the Peruc – Korycany Formation to the upper part of the middle Genus: Brachyphyllum Brongniart 1828 Cenomanian. 1825 Thuites Sternberg, p. 38 [pro parte]. 1870 Echinostrobus Schimper, p. 331. METHODS Type. 1828 Brachyphyllum mamillare Brong- Pieces of carbonised material were carefully picked niart, p. 109. from twig compressions with a preparation needle and treated for cuticle analysis. Carbonized material The form genus, which I understand in the obtained by needle sampling was cleaned by treatment sense of the emendations by Kendall (1947) in HF. After cleaning, it was ready for a bleaching and Harris (1969), is characterized by conifer- procedure that included maceration with Schulze’s ous twigs bearing leaves that have along their reagent: HNO3 + KClO3, neutralisation in water, and treatment in a low concentration solution of KOH, total length leaf cushions that are shorter which was used for washing out the oxidized coal than the width of the leaf cushions. Species of matter. The time for oxidation was about 15 minutes. this form genus belong to the families Chei- After chemical treatment, cuticles were washed in rolepidiaceae (Harris 1979, Watson 1988), water in Petri dishes. Some preparations were stained by a 2% safranine solution in water. Cuticles for light Araucariaceae (Harris 1979) and Taxodiaceae microscopy were embedded in glycerine framed by as pointed out by Clement-Westerhof and Van Noyere framing cement. Usually, preparations with Konijnenburg-Van Cittert (1991). needles under a binocular microscope were necessary to separate lower (abaxial) and upper (adaxial) cuticles and to adjust them properly on the preparation Brachyphyllum squammosum (Velenovský) glass before covering. In this phase of the work, resin Palibin bodies and other remains of mesophyllous tissues were Pl. 1, fi gs 1–4, Pl. 2, fi gs 1–6 isolated and removed. Cuticles prepared for SEM observations were treated in the same way in the Schulze’s reagent, and then were Basionym. Echinostrobus squammosus washed in distilled water. Before drying, cuticles were Velenovský 1885, p. 16, pl. 6, fi gs 3, 6–8. 27

1889 Echinostrobus squammosus Velenovský, (ex 55 mm long. It is covered by helically arranged parte), p. 9, pl. 1, fi gs 13, 14, (non fi gs 16–19); pl. rhomboidal scale-like leaves (4 × 4 mm). Its 2, fi g. 1, (non fi g. 2). branchlets are short, not exceeding 10 mm. 1901 Echinostrobus squammosus Velenovský; Bayer in Frič & Bayer, p. 106, text-fi g. 61, 1, 2 (non The most complete twig, although poorly text-fi g. 61, 3). preserved, is specimen F 00649 and its coun- 1903 Echinostrobus squammosus Velenovský; Bayer terpart F 00650. It represents a 140 mm long in Frič & Bayer, p. 105, text-fi g. 61, 1, 2 (non main axis branched in three orders (Velen- text-fi g. 61, 3). ovský 1889, pl. 2, fi g. 1). The illustration is 1931 Echinostrobus squammosus Velenovský; (ex largely modifi ed and reconstructed. parte) Velenovský & Viniklář, p. 11 (70), (non Specimens known from the Pecínov local- pl. 25, fi gs 2–11). 1937 Brachyphyllum squammosum (Velenovský) Pali- ity (Kvaček & Dilcher 2000) are leaf compres- bin, p. 177. sions showing details of the micromorphology 1968 Echinostrobus squammosus Velenovský; Němejc, of the cuticle. The best-preserved specimen p. 394, pl. 42, fi g. 1. (F 02504) shows a 100 mm long twig branched 1975 Brachyphyllum squammosum (Velenovský) Pali- in three orders, showing blunt terminal bin; Němejc & Kvaček, p. 24, text-fi gs 3, 4; pl. 3, shoots (Pl. 1, fi g. 3). Rhomboidal leaves (Pl. 1, fi gs 7–10; pl. 15, fi gs 1–6. fi g. 2) are hypostomatic, arranged helically 2000 Brachyphyllum squammosum (Velenovský) Pal- ibin; Kvaček & Dilcher, p. 22, pl. 1, fi g. 2. on the twig, and bear delicate grooves on the abaxial side. Lectotype. Designated herein, No. NM The specimen F 02504 provided most of the F 00267, F 00268 (part and counterpart), Vele- material for cuticle analysis. Areas of adaxial novský 1885, pl. 6, fi g. 3, (herein Pl. 1, fi g. 1). and abaxial leaf cuticles are uneven. The adaxial cuticle is confi ned to the narrow area in the Type locality. Vyšehořovice, 30 km east apex of the leaf only. It shows quadrangular or of Prague. polygonal cells (15–35 × 30–65 μm, Pl. 1, fi g. 4) Type horizon. Peruc Korycany Formation, in rows and a narrow cuticular crista (120 μm Cenomanian, Upper Cretaceous. high, Pl. 1, fi g. 2). Ordinary epidermal cells in rows are often orientated transversely, but Emended diagnosis. Leafy twigs, regu- sometimes obliquely or even parallel to the leaf larly branched in one plane; leaves rhomboi- axis with anticlinal walls bent or straight, 1– dal, helically arranged, scale-like, appressed 4 μm thick. The abaxial cuticle shows stomata to the axis, bearing longitudinal grooves arranged in rows, which converge towards the abaxially; free leaf tip very short. Branches of leaf apex (Pl. 1, fi g. 2). Stomatal rows are quite the last order short with blunt apex. Adaxial regular (Pl. 2, fi g. 1). The non-stomatal areas cuticle showing quadrangular or polygonal are one to four cells wide (Pl. 2, fi g. 2). Ordinary cells in rows, the cells transversely orientated. cells are elongate to isodiametric (10–25 × 20– Abaxial cuticle bearing longitudinal interchan- 65 μm) with straight or bent anticlinal walls. ging stomatal and inter-stomatal rows conver- They are arranged in short rows. Stomata are ging towards the apex. Amphicyclic stomata monocyclic or incompletely dicyclic (Pl. 2, fi gs arranged in rows, orientated transversely or 2, 3). Their orientation is not stable. In the obliquely, rarely longitudinally to the leaf apical and basal regions of the leaf lamina the axis. Stomata slightly sunken, surrounded by stomata are usually orientated obliquely or 4–5(6) subsidiary cells; stomatal apparati with longitudinally to the leaf margin while trans- bilobed polar extensions. External leaf surface versally in the central part. The stomata are smooth. surrounded by 4–5(6) subsidiary cells (20–30 Specimens studied. F 00267, F 00268, × 30–55 μm) forming a circular stomatal appa- F 00649, F 00650, F 00867, F 00868, F 02504, ratus (60–100 μm in diameter, Pl. 2, fi gs 3, 4). F 02515, F 02516, F 02838, F 02839. The remains of guard cells are rarely preserved and form a pore 40–45 μm long (Pl. 2, fi g. 3). Occurrence. Vyšehořovice, Pecínov quarry, Sometimes two nodulous polar extensions are unit 5. preserved projecting through the subsidiary Description. The lectotype, designated cells (Pl. 2, fi g. 3). SEM study of the outer abax- herein (Pl. 1, fi g. 1), is preserved as an impres- ial surface of the leaf has documented a rather sion. It represents a part of a branched shoot smooth surface with numerous perforations 28 representing stomatal pits (30–35 μm) without leaves and stomata in rows. It differs from any rim (Pl. 2, fi gs 5, 6). B. squammosum in the presence of the free part of the leaf tip, which is well pronounced. DISCUSSION It is also better preserved – petrifi ed, showing a fl attened wood cylinder. Kunzmann et Brachyphyllum squammosum is a unique al. (2004) described Brachyphyllum obesum taxon known thus far only from the Bohemian Heer sensu Duarte from the Early Cretaceous Cretaceous. Besides the Cenomanian, it was of the Crato Formation in Brazil, which is also also recorded in the South Bohemian Senon- similar to B. squammosum. It has a similar ian (Němejc & Kvaček 1975). This material, cuticle pattern showing stomata in rows, but from the locality of Řídká Blana, agrees in its leaves are amphistomatic. macromorphology with the type material and Brachyphyllum sp. 2 from the Maastrich- also with the material from Pecínov, includ- tian of the Netherlands (Kunrade Region, ing the presence of hypostomatic leaves. It van der Ham & van Konijnenburg-van Cit- differs slightly in cuticle structure. Stomata tert 2003), differs from B. squammosum in of the Cenomanian specimens are not as having amphistomatic leaves and stomata strictly transversely orientated to the leaf randomly distributed only locally arranged margin. This character is considered here in rows. Brachyphyllum patens (Miquel) van as an intraspecifi c variation. Brachyphyllum der Ham et al. from the Maastrichtian of the squammosum described from the Campanian Netherlands and Belgium differs from B. squa- of North Carolina (Raubeson & Gensel 1991) mmosum in having amphistomatic leaves and is very similar in macromorphology, but differs randomly oriented stomata (van der Ham et slightly from B. squammosum of the Czech al. 2003). Brachyphyllum punctatum Miquel Republic in cuticle pattern, having a lower from the English and German Wealden differs density and strictly perpendicular orientation from B. squammosum in having a thick cuti- of stomata in rows. cle with stomata on the abaxial side that are Echinostrobus minor described by Velen- deeply sunken in stomatal tubes (Watson et al. ovský (1889), is a fragment of a conifer twig 1987, 1988). Brachyphyllum tigrense Traverso impression of Brachyphyllum type. Its leaves (1966) from the Lower Cretaceous of Argen- are much smaller, showing pointed apices. It tina differs from B. squammosum in having probably represents a different taxon. Without stomata irregularly distributed. information about its cuticle, comparison with Cuticles of Jurassic members of the genus B. squammosum is diffi cult. (e.g., B. mamillare Lindley & Hutton, B. crucis Most of the Cretaceous species of Brachy- Kendall, B. ardenicum Harris) published by phyllum are preserved as leaf impressions Kendall (1947), Harris (1979) and Orlovskaya or their cuticles have not yet been studied: (1971 – B. mamillareforme Orlovskaja) differ B. obesiforme Saporta, B. confusum Saporta from B. squammosum in having amphistomatic (Cretaceous of Portugal, Teixeira 1948), leaves with stomata distributed irregularly or B. crassicaule Fontaine, (US Cretaceous, in short or irregular rows. Within Jurassic Potomac F., Fontaine 1889), B. crassum species, the macromorphology of B. squammo- Lesquereux = B. macrocarpum Newberry nom. sum resembles most B. mamillare associated inval. (US Cretaceous, Dakota F., Lesquereux with Araucaria philipsii (Carruthers) Schim- 1892), B. araxenum Palibin (Cretaceous of per (Harris 1979). Dalaragez, Armenia, Palibin 1937). Brachy- The arrangement of stomata (orientated phyllum sp. from the Cretaceous of Aachen transversely to the leaf margin) of Brachy- (Stockmans 1946), is very similar to B. squam- phyllum squammosum is more similar to the mosum in macromorphology (as far as it is pos- genus Agathis, particularly in the presence of sible to recognize from the simplifi ed illustra- stomatal polar extensions. However, Velen- tion), but it is an impression specimen. Ohana ovský and Viniklář (1931) state that sterile and Kimura (1993) described material labelled twigs of Brachyphyllum squammosum were as Brachyphyllum vulgare (Stopes & Fujii) found attached to taxodiaceous cones at the Jeffrey from the Upper Cretaceous of Japan. locality of Jevíčko. Unfortunately, their mate- It agrees in many characters with B. squam- rial was not fi gured and was not available mosum, particularly in having hypostomatic for the present study. At present there is no 29 clear evidence for the taxodiaceous affi nity of BRONGNIART A.T. 1828. Prodrome d’une histoire Brachyphyllum squammosum. On the other des végétaux fossiles. F.G. Levrault, Paris. hand, there is no robust support for its affi ni- ČECH S., KLEIN V., KŘÍŽ J. & VALEČKA J. 1980. ties with the family Araucariaceae. Revision of the Upper Cretaceous stratigraphy of the Bohemian Cretaceous Basin. Věst. Ústř. Úst. In association with sterile foliage of Bra- Geol., 55(5): 277–296. chyphyllum squammosum, Velenovský (1885) CLEMENT-WESTERHOF J.A. & van KONIJNEN- described a cone-like reproductive structure BURG-van CITTERT J.H.A. 1991. Hirmeriella consisting of bilobed “scales”. The structure muensteri: New data on the fertile organs leading was classifi ed by Bayer (1914) as Strobilostro- to a revised concept of the Cheirolepidiaceae. Rev. bus nom. nud. and by Velenovský and Viniklář of Palaeob. Palynol., 68: 147–179. as Diplostrobus stupeckyanus Velenovský EKLUND H. & KVAČEK J. 1998. Lauraceous infl o- & Viniklář (1931). Although in both later pub- rescences and fl owers from the Cenomanian of lications the authors realised the independent Bohemia (Czech Republic, Central Europe). Intern. Jour. Plant Sci., 159: 668–686. status of a sterile twig (B. squammosum) and the reproductive structure, the problem of the FONTAINE W.M. 1889. The Potomac or Younger Mesozoic Flora. U.S. Geol. Surv. Monographs, 15: systematic affi nity of the reproductive organ 1–377. consisting of bilobed structures remained FRIČ A. & BAYER E. 1901. Studien im Gebiete der unresolved. In both cases Bayer (1914) and Böhmischen Kreideformation. Perucer Schichten. Velenovský and Viniklář (1931) interpreted Archiv der naturwisschenschaftlichen Landes- the structure as a problematical conifer cone. durchforschung von Böhmen, 11(2): 1–180. However, new studies (Eklund & Kvaček 1998, FRIČ A. & BAYER E. 1903. Studie v oboru křídového J. Kvaček unpublished data) indicate that útvaru českého. Perucké vrstvy. Archiv pro Diplostrobus is an angiosperm infl orescence. přírodovědecké prozkoumání Čech, 11(2): 1–179. As in the case of Mauldinia (Lauraceae), the HARRIS T.M. 1969. Naming of fossil conifer: 243–252. bilobed structures of Diplostrobus probably In: Santarau H., Ghosh A.K., Roy S.K., Chanda S. represent cladodia of an infl orescence of a lau- & Chaudhuri S.K. (eds) J. Sen Memorial Volume. roid angiosperm. Botanical Society of Bengal, Calcutta. HARRIS T.M. 1979. The Yorkshire Jurassic flora, Ecological remarks. Brachyphyllum 5. Trust. British Mus. (Nat. Hist.), London. squammosum (Velenovský) Palibin is typically KENDALL M.W. 1947. On five species of Brachyphyl- found as small fragments. In Pecínov it occurs lum from the Jurassic of Yorkshire and Wiltshire. in a Sphenolepis assemblage together with Ann. Magaz. Nat. Hist., Ser. 11, 14: 225–251. Sphenolepis pecinovensis (Kvaček 1997) and KUNZMANN L., MOHR B.A.R. & BERNARDES- Cunninghamites lignitum (Kvaček 2000). The DE-OLIVEIRA M.E.C. 2004. Gymnosperms from presence of other mesophyte taxa (Gnetales, the Lower Cretaceous Crato Formation (Brazil). Nilssoniopteris pecinovensis Kvaček 1995) and I. Araucariaceae and Lindleycladus (Inceratae sedis). Mitteil. Mus. Naturk. Berlin, Geowiss. its habit suggest that Brachyphyllum squam- Reihe, 7: 155–174. mosum was probably a member of “upland” vegetation. KVAČEK J. 1995. Cycadales and Bennettitales leaf compressions of the Bohemian Cenomanian, Cen- tral Europe. Rev. Palaeobot. Palynol., 84: 389–412. ACKNOWLEDGEMENTS KVAČEK J. 1997. Sphenolepis pecinovensis sp. nov. a new taxodiaceous conifer from the Bohemian This work was supported by a grant of the Min- Cenomanian. Meded. van ‘s Rijks Geol. Dienst. istry of Culture of the Czech Republic (No. MK Haarlem, 58: 121–128. 00002327201). I thank to Jana Čejková and my wife Helena who facilitated SEM analysis and Arden Bash- KVAČEK J. 2000. Two conifers (Taxodiaceae.) of the forth for English editing. The paper profi ted from the Bohemian Cenomanian, Czech Republic, Central constructive comments of two anonymous referees. Europe. Acta Palaeobot., Suppl. 2: 129–151. KVAČEK J. & DILCHER D. 2000. Comparison of Cenomanian fl oras from Western Interior North REFERENCES America and Central Europe. Acta Univ. Carol., Geol., 44: 17–38. LESQUEREUX L. 1892. The Flora of the Dakota BAYER E. 1914. Fytopaleontologické příspěvky ku Group. U.S Geol. Surv., Monographs, 17: 1–256. poznání českých křídových vrstev peruckých. Archiv pro přírodovědecké prozkoumání Čech, NĚMEJC F. 1968. 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NĚMEJC F. & KVAČEK Z. 1975. Senonian plant high-frequency sea level change in fl uvial estua- macro fossils from the region of Zliv and Hluboká rine succession: Cenomanian palaeovalley fi ll, (near České Budějovice) in South Bohemia. Uni- Bohemian Cretaceous Basin: 247–268. In: Howell versita Karlova, Praha. J.A. & Aitken J.-F. (eds) High-resolution Sequence Stratigraphy innovation and applications. Geol. OHANA T. & KIMURA T. 1993. Permineralized Soc. Spec. Publ. 104, London. Brachyphyllum leafy branches from the Upper Yezo Group (Coniacian-Santonian) Hokkaido, ULIČNÝ D., KVAČEK J., SVOBODOVÁ M. & ŠPI- Japan. Bull. National Sci. Mus., Series C (Geol. ČÁKOVÁ L. 1997. High-frequence sea-level fluc- & Paleont.), 19(2): 41–54. tuations and plant habitats in Cenomanian fl uvial to estuarine successions: Pecínov quarry, Bohe- ORLOVSKAYA E.R. 1971. Predstaviteli Brachyphyl- mia. Palaeogeogr. Palaeoclimat. Palaeoecol., 136: lum i Pagiophyllum v pozdney yure khrebta Kara- 165–197. tau (Representatives of Brachyphyllum and Pagio- phyllum in the Late Jurassic of Karatau Ridge). Van der HAM R.W.J.M. & van KONIJNENBURG- Materialy po istorii fauny i fl ory Kazakhstana, 5: van CITTERT J.H.A. 2003. Rare conifers from the 66–77. (in Russian). type area of the Maastrichtian (Upper Cretaceous, southeast Netherlands). Scripta Geologica, 126: PACLTOVÁ B. 1977. Cretaceous angiosperms of Bohe- 111–119. mia – Central Europe. Bot. Rev., 43(1): 128–142. Van der HAM R.W.J.M & van KONIJNENBURG- PALIBIN I.V. 1937. Melovaya flora Daralageza (The van CITTERT J.H.A., DORTANGS R.W., Cretaceous fl ora of the Daralagez). Acta Inst. Bot. HERNGREEN J. & van der BURGH J. 2003. Acad. Sci. URSS, Ser. 1, 4: 171–197. (in Russian). Brachyphyllum patens (Miquel) comb. nov. (Chei- RAUBESON L.A. & GENSEL P.G. 1991. Upper Cre- rolepidiaceae?): remarkable conifer foliage from taceous conifer leaf fossils from the Black Creek the Maastrichtian type area (Late Cretaceous, NE Formation with an assessment of affi nities using Belgium, SE Netherlands). Rev. Palaeobot. Paly- principal components analysis. Bot. Gaz., 152: nol., 127: 77–97. 380–391. VELENOVSKÝ J. 1885. Die Gymnospermen der Böh- SCHIMPER W.P. 1870. Traité de paléontologie mischen Kreideformation. E. Greger, Prag. végétale ou la fl ore du monde primitif dans ses VELENOVSKÝ J. 1889. Květena Českého cenomanu. rapports avec les formations géologique et la fl ore Rozpravy Královské České Společnosti Nauk, 7(3): du monde actuel, vol. 2, part 1. J. B. Bailliére et 1–75. fi ls, Paris. VELENOVSKÝ J. & VINIKLÁŘ L. 1931. Flora Creta- STERNBERG K.M. 1825. Versuch einer geognostisch- cea Bohemiae 4. Stát. Geol. Ústav Československé botanischen Darstellung der Flora der Vorwelt. Republiky, Praha. Ernst Brenck’s Wittwe, Regensburg. WATSON J. 1988. The Cheirolepidiaceae: 382–447. STOCKMANS F. 1946. Végétaux de l’assise des sables In: Beck, C.B. (ed.) Origin and evolution of gymno- d’Aix-La-Chapelle récoltés en Belgique (Sénonien sperms. Columbia University Press, New York. inférieur). Mém. Mus. Royal Hist.Natur. Belgique, 105: 1–51. WATSON J., FISCHER H. & HALL N.A. 1987. A new species of Brachyphyllum from the English TEIXEIRA C. 1948. Flora Mesozóica Portugesa, Wealden Formation. Rev. Palaeobot. Palynol., 51: I parte. Serviços geológicos de Portugal, Lisboa. 169–187. TRAVERSO N.E. 1966. Brachyphyllum tigrense, WATSON J., FISCHER H. & HALL N.A. 1988. The nueva conífera de la Formación Baqueró, Cretácico holotype of the Wealden conifer Brachyphyllum de Santa Cruz. Ameghiniana, 4(6): 189–194. punctatum Michel. Palaeontology, 31(4): 1029– ULIČNÝ D. & ŠPIČÁKOVÁ L. 1996. Response to 1031. PLATES 32

Plate 1

Brachyphyllum squammosum (Velenovský) Palibin

1. Lectotype, shoot, × 2, Vyšehořovice; F 267 2. LM of apical part of leaf, × 40, Pecínov; F 2504b 3. Branched shoot, × 1, Pecínov; F 2504 4. LM of adaxial cuticle, × 200, Pecínov; F 2504c Plate 1 33

J. Kvaček Acta Palaeobot. 47(1) 34

Plate 2

Brachyphyllum squammosum (Velenovský) Palibin

1. LM of abaxial cuticle, × 40, Pecínov; F 2504b 2. LM of abaxial cuticle, × 200, Pecínov; F 2504c, 3. LM of abaxial cuticle, stoma, × 900, Pecínov; F 2504c 4. SEM of abaxial cuticle, inner surface, × 300, Pecínov; F 2504d (7457) 5. SEM of abaxial cuticle, outer surface, stoma, × 1000, Pecínov; F 2504d (7480) 6. SEM of abaxial cuticle, outer surface, stomatal rows, × 300, Pecínov; F 2504d (7477) Plate 2 35

J. Kvaček Acta Palaeobot. 47(1)