Cenomanian Florule of Nammoura, Lebanon

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Cretaceous Research (2000) 21, 785–799 doi:10.1006/cres.2000.0229, available online at http://www.idealibrary.com on Cenomanian ﬂorule of Nammoura, Lebanon

*Valentin Krassilov and †Flavio Bacchia

*Paleontological Institute, 1123 Profsoyusnaya St., Moscow 117641, Russia †Stoneage S. R. L., Via Torino, 15, 34123 Trieste, Italy

Revised manuscript accepted 8 August 2000

The mid-Cenomanian fish beds of Nammoura, Mont-Liban district, Lebanon contain a diverse fauna of aquatic and terrestrial vertebrates, a few crustaceans and moderately well-preserved plant remains of which a single species, Sapindopsis anhouryi, was previously described by Dilcher & Basson (1990). We add 11 species of ferns, gymnosperms and angiosperms of which Nammouria cretacea gen. et sp. nov, Nupharanthus cretacea gen. et sp. nov., Sapindopsis libanensis sp. nov. and Nammourophyllum altingioides gen. et sp. nov. are new taxa. The florule differs markedly from both Early Cretaceous and Turonian plant assemblages of the Middle East, thereby representing a distinct stage of the regional floristic evolution. Its phytogeographic affinities are with contemporaneous floras of North America, Central Europe and the Crimea. A combination of such features as xeromorphism, the prevalence of compound leaves, and the presence of deciduous angiosperm components and gymnosperms may indicate climatic conditions similar to those of the present day Mediterranean. 2000 Academic Press

K W: aquatic ferns; early angiosperms; Middle East; Cretaceous; palaeoclimates.

1. Introduction the Milan Museo Civico di Storia Naturale and in private collections. The material described in this paper comes from two The terrestrial plants are preserved as impressions quarries in the Al Gabour Valley near the village of with fragmentary compression films. Previously, a Nammoura, Mont-Liban district, about 20 km north- single plant species, Sapindopsis anhouryi, was de- east of Beirut and about 10 km south-southeast of scribed from Nammoura by Dilcher & Basson (1990). Hjoula, a well-known Cenomanian fish locality. The We add 11 forms of ferns, gymnosperms and quarries cut a sequence of Cretaceous platform car- angiosperms. Natalia P. Maslova (Palaeontological bonates with a fossiliferous horizon, the fish beds, near Institute, Moscow) contributed to the recognition and its base. The fish beds incorporate the same facies as description of the new species Sapindopsis lebanensis those represented at Haqel and Hjoula (Hu¨ckel, and Nammourophyllum altingioides. The fossil plant 1970). They consist of alternating carbonaceous collection is presently housed in the Paleontological mudstones and wackestones followed up-section by Institute, Moscow, catalogue no. 4799. greenish or bluish-grey, thinly laminated limestones with chert nodules, altogether 15–30 m thick. The fish beds are overlain by a rudist bank, the foraminiferal 2. Systematic palaeontology assemblage of which contains a characteristic Ceno- manian species Rhapydionina laurinensis Saint-Marc. Pteridophytes In addition to the diverse medium-sized to large fish remains, including a large shark tooth, the fossiliferous Pteris sp. horizon has yielded a few small crustaceans, coprolites Figure 1C–F and fecal pellets a few millimeters in diameter. There are also dismembered skeletons of aquatic and terres- Description. Three fragments of pinnate leaves 50 mm trial reptiles, including a forearm of a large theropod long, with a thin, longitudinally striate, rachis distally dinosaur and a few bird feathers. The holotype of forking at an acute angle. The pinnae are alternate, the recently described dolichosaur Aphanizocnemus well spaced on the main rachis, more crowded on the libanensis Dal Sasso & Pinna (1997) supposedly came distal arms, arising at about 60–80, lanceolate, up to from this locality. The faunal remains are housed in 7 mm long, variably dissected, with a pair or two pairs

0195–6671/00/060785+15 $35.00/0 2000 Academic Press 786 V. Krassilov and F. Bacchia

Figure 1. Cenomanian plants of Nammoura, Lebanon. A, B, Pseudolarix sp., conifer needle, no. 4799/4,1 and 15. C–F, Pteris sp., fern leaf fragments, no. 4799/15, 1; pinna enlarged to show marginal coensori, 8; and cuticle with hair bases, SEM, 400. G–I, Pseudotorellia sp., a ginkgophyte leaf, no. 4799/1, 1 and enlarged, 7, to show resin canals (dark slightly sinuous lines); note a transverse insect mine (arrows). Cenomanian florule of Nammoura, Lebanon 787 of proximal pinnules, the latter distally slightly incised order, bearing thick, elliptical to spindle-shaped or entire. The pinnules are anadromous or catadro- hirsute bodies (sporocarps?) at each branching node. mous, elliptical to rounded, in the distal pinnae re- Ultimate divisions of the marginal pinnae might also duced to lobes. A typical condition of the distal pinnae terminate in sterile pinnules, but these are either shed is a short lanceolate blade with a single abaxial lobe or poorly preserved. and a shallow adaxial incision. The venation is pin- The sterile pinnule is broadly lanceolate, 2 mm nate, with 2–3 pairs of lateral branches in the entire wide, with six diverging, sporadically anastomosing part of the pinna, 1–2 branches in the basal pinnules. veins (Figure 4B). The hirsute bodies arise singly or in The fertile pinnae show linear marginal coenosori pairs at an acute angle to the rachis. They are sessile, 0.3 mm wide. Our attempts at obtaining spores failed, slightly decurrent, elliptical to spindle-shaped, acumi- but the pinnae yielded thin cuticle with rounded- nate, the proximal ones relatively large, about 5 mm polygonal cells (Figure 1F). long, 2 mm wide, distally decreasing to 31mm. Their surface is pitted, with long hairs leaving faint Remarks. The pinnule morphology and the position of impressions along the margins (Figure 4A, B). Under coenosori are rather typical of the extant genus Pteris an SEM, the ferruginous impressions show reticulate (Pteridaceae). Forking leaves occasionally occur in the spherical structures about 90–100 m in diameter that closely related family Vittariaceae (Copeland, 1945). appear to be megaspores (Figure 4C), but their No comparable forms are currently known among preservation is too poor for a definite interpretation. Cretaceous ferns. Remarks. The leaf has a distinctive morphology that is Nammouria Krassilov gen. nov. not matched by any known Mesozoic plant. It is, therefore, deserving a status of a new morphological Derivation of name. From the Nammoura locality. genus. In the lack of well-preserved structural details its taxonomic assignment is admittedly ambiguous. A Type species. Nammouria gracilis sp. nov. comparison with aquatic ferns is based on the morphology of both the sterile pinnule and hirsute bodies Diagnosis. Leaf stalked, dichopodially divided, with that resemble, respectively, the floating leaf segments the pinnae repeatedly forked. Sterile pinnules terminal and sporocarpia of extant Marsilea and Regnellidium, on pinna rachis, broadly lanceolate, with flabellate as well as those of fossil Rhodeites and Hydropteris venation. Elliptical to spindle-shaped hirsute bodies (sporocarps?) are born singly or in pairs at each (Chitaley & Paradkar, 1972; Rothwell & Stockey, branching node. 1994). In these ferns sporocarpia typically attach at the branching nodes of the rhizome. Dichopodial Nammouria gracilis Krassilov, sp. nov. branching occurs in submerged leaves of the Salvin- Figures 2C, D, 3, 4A–C iaceae, but has not been recorded hitherto in the floating or emergent leaves. Nammouria might repre- Holotype. Palaeontological Institute, Moscow, Cata- sent an extinct line of initial hydropterid radiation that logue no. 4799-10, Nammoura locality, Lebanon. took place in the Early to mid-Cretaceous (Krassilov & Golovneva, 1999). Diagnosis. As for the genus. Gymnosperms Description. The species is based on the part and counterpart of a stalked, dichopodially divided leaf Pseudotorellia sp. (with the exterior segments overtopping the interior Figure 1G–I ones). The stalk is straight, 20 mm long, 1 mm thick, longitudinally striate, distally divided into three Description. A lanceolate leaf, 677 mm, bluntly pinnae that spread in one plane, apparently arising pointed, with margins parallel for most of the length, from one point, but actually produced by two dichoto- gradually converging to the base which shows a thick- mies rapidly following one another (Figure 2C, D). ened abscission scar (Figure 1I). The veins diverge The median pinna is relatively short, bearing a termi- from the base, subparallel above, seven in the middle nal sterile pinnule. The marginal pinnae diverge at part. Irregularly twisted resin ducts occur between the about 60. Their rachises are less than 0.5 mm thick, veins. The blade is traversed by a leaf mine of the type curved, with 2–3 longitudinal furrows, repeatedly described by Labandeira et al. (1994, Figure 1I–L)on forked at intervals 7, 4 and 2 mm in distally decreasing Densinervum sp. from the Dakota Formation. 788 V. Krassilov and F. Bacchia

Figure 2. Cenomanian plants of Nammoura, Lebanon. A, B, Phragmites sp., monocot leaf, no. 4799/13, 1 and detail of venation (B, orientated at 90 to A), SEM, 50. C, D, Nammouria gracilis gen. et sp. nov., aquatic fern (?), holotype no. 4799/10a, dichopodially divided leaf bearing sterile pinule (arrow) and sporocarps (?), 1 and 3.5. Cenomanian ﬂorule of Nammoura, Lebanon 789

Figure 3. Nammouria gracilis gen. et sp. nov., aquatic fern (?) from the Cenomanian of Nammoura, Lebanon: sketch drawing of holotype, 1.5.

Remarks. Of the pseudotorellias with resin ducts, Remarks. This leaf might belong to a deciduous P. linkii (Ro¨mer) Watson is similar but considerably conifer with Larix-Pseudolarix aﬃnities. It is similar smaller (Watson & Harrison, 1998). to the extant Pseudolarix kemferi Gord., the Paleogene P. nipponica Kimura & Horiuchi (1978) and the Early Pseudolarix sp. Cretaceous P. erensis (Krassilov, 1982). In this last Figure 1A, B species, the leaf dimensions and the width ratio of the costal and stomatal zones exactly match the Lebanon Description. A linear-oblanceolate needle-leaf, 25 mm species. long, 2 mm wide. The leaf is widest below the apical constriction from where it gradually tapers towards Monocotyledons the base, lacking a distinct petiole, but with the base constricted and twisted, showing a straight abscission ‘Phragmites’ sp. line. The apex forms a short, blunt point. The leaf Figure 2A, B surface shows a median costal zone 1 mm wide, wedging out a little below the apex, and two slightly Description. A ribbon-shaped leaf, more than 155 mm depressed submarginal stomatal zones, each 0.3 mm long (apex lacking), 13 mm wide at the base, ex- wide. panded to 16 mm slightly below the middle, gradually 790 V. Krassilov and F. Bacchia

Figure 4. Nammouria gracilis gen. et sp. nov., aquatic fern (?) from the Cenomanian of Nammoura, Lebanon: holotype, no. 4799/13. A, B, hirsute bodies (sporocarps?) showing pitted surface and hairs impressed along the margins (arrow); part of a sterile leaf is seen in B, left, 10. C, SEM of a hirsute body showing a spherical reticulate structure (megaspore?), 500.

tapering toward the apex. The base is slightly concave, Heer: Hollick, 1936), but their affinities to this genus thickened along the abscission line. The veins are and to gramineous monocots have yet to be proved. straight, parallel, 1–0.7 mm apart, with the interstitial veins traceable for a short distance. Epidermal cells Dicotyledons are arranged in files parallel to the veins (Figure 2B). Nupharanthus Krassilov gen. nov. Remarks. This leaf is of the same type as the widespread Late Cretaceous and Cenozoic remains tradi- Derivation of name. From extant genus Nuphar tionally assigned to Phragmites (e.g. Pragmites alaskana (Nymphaeaceae), and anthus, flower. Cenomanian florule of Nammoura, Lebanon 791

Type species. Nupharanthus cretacea sp. nov. (Manchester, 1992), but the sepals are connate, with a distinct reticulate venation. Diagnosis. Flower solitary, perfect actinomorphic. Five sepals, free, imbricate, with subparallel venation. Sapindopsis ahhouryi Dilcher & Basson Petals smaller than sepals in more than one circle, Figure 7 heteromorphous. About ten stamens, spirocyclic, leaving elongate scars with pits of vascular bundles. 1990 Sapindopsis anhouryi Dilcher & Basson, 1990, Stigmatic disk faintly striated radially. p. 540, figs 1–16. Description. A compound leaf with four alternate leaf- Nupharanthus cretacea Krassilov, sp. nov. lets and a sheathing stipule at the base. The leaflets are Figures 5A, B, 6 linear-lanceolate, about 65–807 mm, slightly asymmetrical at the base, decurrent, with stout midrib, thin Derivation of name. From the Cretaceous. secondaries and a prominent intramarginal vein. The stipule is ovate, 156 mm, with subparallel veins Holotype. Palaeontological Institute, Moscow, Cata- converging to the apex. logue no. 4799-9, Nammoura locality, Lebanon. Remarks. This species differs from the contempor- Diagnosis. As for the genus. aneous North American Sapindopsis (Huang & Dilcher, 1994) by its better-developed intramarginal Description. The flower is preserved as a pale yellowish vein, which gives the venation a myrtaceous aspect. In impression showing a radially symmetrical calyx leaf shape and venation it is also comparable with 25 mm in diameter, with five sepals spreading in the Halyserites reichii Sternberg from the Cenomanian of bedding plane, slightly imbricate proximally, distally the Bohemian Massif (Knobloch, 1978). spreading, with broadly rounded apices. The sepal venation is clearly marked, consisting of thick subpar- Sapindopsis lebanensis Krassilov & Maslova, sp. nov. allel veins, with indistinct cross veins. Intersecting Figures 8, 9A–D vein patterns are seen in the marginal overlap areas, indicating the membranous texture of the sepals. Holotype. Palaeontological Institute, Moscow, Cata- The petals are of variable shapes and dimensions logue no. 4799-11, Nammoura locality, Lebanon. ranging from semicircular to lingulate, 3.5–5mm long. They are either distinctly or faintly impressed Diagnosis. Leaf lobate-palmate, stipulate, with a basal upon the sepals, apparently arranged at different levels leaflet distinct, the rest increasingly fused over the leaf. in a spirocyclic series. The petal venation is thin, Leaflets linear-lanceolate, about 404 mm, acutely digitate, with oblique anastomoses. pointed. Midrib prominent, secondaries thin, faintly The stamens are shed, but about ten scars are marked, eucamptodromous, joining intramarginal conspicuous in the peripheral zone of the receptacle vein. Stipule sheathing ovate-lanceolate, parallel- (Figure 5A). The scars are arranged in an irregular veined, with free part triangular, 1 mm long. Leaf spirocyclic series, and are elongate, slightly curved, compression thick, with stomatal pits distinct at low with rounded pits apparently left by vascular bundles. magnification, and with transverse striation reflecting The central part of the receptacle is occupied by a disk pallisade mesophyll structure. Leaf blade hyposto- 2 mm in diameter, with a slightly undulating margin. matic, with hair bases and large spherical glands The disk surface shows faint radial striations. (Figure 9B) on both sides. Secondary veins scarcely reflected in the epidermal cell patterns. Ordinary Remarks. Distinguishing features of the flower are its epidermal cells polygonal, irregularly disposed or pentamerous actinomorphic structure, distinct sepals, radiating from a larger, rounded cell. Stomata scat- and spirocyclic arrangement of small petals and tered, irregularly orientated, anomocytic to weakly stamens which, judging by the scars, might have been cyclocytic, with typically five trapezoid subsidiary cells laminar. These features indicate affinities with the that are slightly sunken below the level of ordinary Nymphaeaceae, in particular with the extant genus cells. Some guard cells show long, T-shaped polar Nuphar, which has thin membranous sepals and re- thickenings. Hair bases small, elliptical, encircled by a duced petals (Mosley, 1972). No fossil genus matches variable number (up to ten) epidermal cells. the description. The Palaeogene Florissantia, assigned to the Malvales, is similar in general shape, dimen- Remarks. This species belongs to a xeromorphic group sions and the relative development of sepals and petals of Sapindopsis with narrow leaflets, a thick, probably 792 V. Krassilov and F. Bacchia

Figure 5. Cenomanian plants of Nammoura, Lebanon. A, B, Nupharanthus cretacea gen. et sp. nov., nymphaeaceous flower, holotype no. 4799/9, showing large, imbricate sepals and smaller, indistinctly marked petals (arrows), 2.5 and 10; insertion, top right corner, shows central part differently illuminated with distinct scars of stamens. C, Aryskumia sp. cf. A. zelkovifolia Shilin, serrate angiosperm leaf with a short twisted petiole, no. 4799/12, 1. D, E, Nammourophyllum altingioides gen. et sp. nov., serrate angiosperm leaf, holotype no. 4799/11, 1, and detail of marginal serration, 7. Cenomanian florule of Nammoura, Lebanon 793

leaflets and well-marked venation. Sapindopsis lebanensis differs from the North American species in its relatively smaller dimensions, better developed intramarginal vein and stipule characters. Their cuticular structures are similar (see Upchurch, 1984), although in the Lebanese species the lateral to polar differenti- ation of subsidiary cells is less distinct. Proteophyllum laminarium Velenovsky´ from the Cenomanian of Bohemia has a slightly broader lamina which is similar in its coriaceous texture, indistinct secondaries and cuticular structure (Neˇmejc & Kvacˇek, 1975). Proteophyllum conventionally includes simple leaves, some of which, however, appear as Sapindopsis-type Figure 6. Nupharanthus cretacea gen. et sp. nov., leaflets. nymphaeaceous flower from the Cenomanian of Nammoura, Lebanon: sketch of holotype, 2.5. Nammourophyllum Maslova & Krassilov gen. nov. coriaceous leaf blade, indistinct venation, amphisto- Derivation of name. From Nammoura locality. matic epidermis and sunken stomata. In contrast, Sapindopsis anhouryi Dilcher & Basson represents a Type species. Nammourophyllum altingioides Maslova & relatively mesomorphic type of leaf with much broader Krassilov sp. nov.

Figure 7. Sapindopsis anhouryi Dilcher & Basson, 1990, compound stipulate angiosperm leaf, no. 4799/14, from the Cenomanian of Nammoura, Lebanon, 1. 794 V. Krassilov and F. Bacchia

Derivation of name. From extant genus Altingia (Hamamelidales).

Holotype. Palaeontological Institute, Moscow, Cata- logue no. 4799-7, Nammoura locality, Lebanon.

Diagnosis. As for the genus.

Description. A single complete leaf with a blade 32 mm long, reaching 15.5 mm in maximum width slightly above the middle. The petiole is 13 mm long, 1 mm thick, expanded to 1.5 mm at the abscission line. The leaf base is broadly cuneate, asymmetrical, with the halves of the blade slightly displaced against each other. The apex is a short point, appearing as a larger marginal tooth. The margin is serrate from the base up. The serration is somewhat irregular, with the teeth tending to increase distally. Larger teeth in the distal part of the blade bear a small additional serration (Figure 5E). The marginal glands are conspicuous in the larger teeth (Figure 10). The midrib starts as a direct continuation of the petiole, gradually attenuating to the apex. Both the petiole and midrib are longitudinally striate. The secondaries are thin and indistinct, arising at variable angles and not quite parallel to each other. The lower secondaries are short, not ascending above the basal quarter, recurved. The next pair ascends at a more acute angle giving oﬀ a few basiscopic branches. These and subsequent secondaries loop irregularly well before the margin. Short branches ascending from the loops terminate in the sinuses of marginal serrations.

Remarks. This leaf represents a distinct morphotype with no close allies in Cretaceous leaf ﬂoras. Crassi- denticulatum Upchurch & Dilcher, 1990 is similar in the glandular serration and irregular brochidodro- Figure 8. Sapindopsis libanensis sp. nov., compound stipu- mous venation, but the marginal teeth are much late angiosperm leaf from the Cenomanian of Nam- smaller and of chloranthoid morphology, while the moura, Lebanon, holotype no. 4799/7, 2. secondaries are far more numerous. Hamamelites Saporta is similar in leaf shape (though more con- Diagnosis. Leaf small, elliptical, long-petiolate, base spicuously asymmetrical in the type species H. clarus slightly asymmetrical, margin irregularly serrate, with Saporta, 1868) and secondary venation, but the rare additional serrations. Larger serrations glandular. petiole is typically short and the marginal teeth are Venation pinnate, brochidodromous. Midrib stout, non-glandular. secondaries thin, irregularly spaced. Intrabasal sec- The marginal features of Nammourophyllum are ondaries short, recurved. Subsequent secondaries rather altingioid, with the secondaries or their derived curved upwards, looping far from the margin, their veins ending in the sinuses. In herbarium material of distal branches terminating in sinuses of the marginal Altingia excelsa we observed rare additional teeth over serration. the typically simple marginal serration. The striate petiole (long in A. gracilis) continuing into the blade as Nammourophyllum altingioides Maslova & Krassilov a similarly striate midrib is also an altingioid feature. sp. nov. However, in modern Altingiaceae the secondaries are Figures 5D, E, 10 far more regular, looping closer to the leaf margin. Cenomanian ﬂorule of Nammoura, Lebanon 795

Figure 9. Sapindopsis libanensis sp. nov., from the Cenomanian of Nammoura, Lebanon, holotype no. 4799/7, SEM micrographs of abaxial cuticle with stoma (A), epidermal gland (B), and hair bases (C, D). 796 V. Krassilov and F. Bacchia

Remarks. In the marginal characters and venation this leaf resembles Aryskumia zelkovifolia, a primitive ulmaceous morphotype, but is somewhat smaller and with a denser serration than the type material from the Santonian of Kazakhstan (Shilin, 1986).

Parvileguminophyllum sp. Figure 11D

Description. A complete leaﬂet, oblanceolate, 3010 mm, gradually tapering to the base, apparently with a short petiole, and with a bluntly pointed apex. The margin is entire, indistinctly undulate, incised by a marginal gall (at left side near the apex in Figure 10D). The midrib extends to the apex. The secondaries are thin, eucamptodromous.

Remarks. The leaf genus Parvileguminophyllum was erected by Herendeen & Dilcher (1990) for compound leaves and small inequilateral leaflets which could be assigned to the Leguminosae. The leaf from Nammoura is considerably broader than the Eocene Figure 10. Nammourophyllum altingioides gen. et sp. species and lacks a mucronate apex, which is nov., serrate angiosperm leaf from the Cenomanian of characteristic of the latter (Call & Dilcher, 1994). Nammoura, Lebanon: sketch drawing of holotype, 2. 3. Discussion The Nammoura florule appears to represent a distinct Platycaryeae gen. et sp. indet. stage of floristic evolution in the Middle East. It is Figure 11A–C fairly different from both the Early Cretaceous and Turonian–Maastrichtian regional floras. No elements Description. An infructescence of bracteate fruits are shared with the widespread Jurassic–Early Creta- 30 mm long, 10 mm wide, dense, with a short, stout ceous Piazopteris-Brachyphyllum assemblages (e.g., El peduncle. The bracts are crowded, showing obliquely Chair et al., 1995; Barale et al., 1997), which suggests spreading acuminate beaks that emerge from the mass a major mid-Cretaceous floristic change. The Nam- of intact fruits (Figure 11B–C). The latter are poorly mouran ferns and gymnosperms are morphologically preserved, some showing relatively broad, mem- modern, with the mid-Mesozoic elements represented branous lateral wings that spread over the bedding by Pseudotorellia alone. The Early Turonian flora of plane. Israel is comparable on account of the putative gramminoid elements (Krassilov & Dobruskina, 1998), but Remarks. The infructescence is of a general juglanda- is dominated by the broad-leaved platanoid morpho- ceous aspect and resembles Platycarya americana from types (Dobruskina, 1996) that are lacking in the the Lower Eocene of North Dakota (Wing & Hickey, Nammoura locality. 1984), but details of the nutlet morphology are The Nammouran angiosperm affinities are with the scarcely discernible. Cenomanian Sapindopsis of North America and their coeval Proteophyllum of Central Europe. Aryskumia Aryskumia sp. cf. A. zelkovifolia Shilin is a modern-looking Paratethyan element previously Figure 5C described from western Kazakhstan and the Crimea (Shilin, 1986; Krassilov, 1984). Description. A poorly reserved leaf with ovate blade, The Nammoura assemblage is too small for a 5020 mm, showing a doubly serrate margin and comprehensive palaeoecological analysis. However, craspedodromous venation in the proximal part of the some preliminary considerations seem justified by the blade. The leaf base is slightly cordate, with a short material. Most importantly, the Cenomanian assem- (4 mm), stout, twisted petiole. blage gives no evidence of a tropical climate. It is Cenomanian florule of Nammoura, Lebanon 797

Figure 11. Cenomanian plants of Nammoura, Lebanon. A–C, Platycaryeae gen. sp., juglandaceous catkin, no. 4799/8, 3, and details of bracts (B) and nuts (N), 10 and 15. D, Parvileguminophyllum sp., legume leaﬂet with a marginal gall (arrow), no. 4799/2, 2.

relatively small-leaved, with a distinct xeromorphic Morphologically, the assemblage hosts two groups component (Pseudotorellia with thick, narrow, resinif- of angiospems: those with compound leaves (Sapin- erous leaves; Sapiondopsis lebanensis with narrowly dopsis, Parvileguminophyllum, the Platycaryeae) and dissected, thick, coriaceous, thickly cutinized, amphi- those with simple serrate leaves (Aryskumia, Nam- stomatic glandular leaves; even the fern leaves are mourophyllum). The compound leaf morphotypes have quite small, with resistant cuticle). In comparison narrow, entire leaflets. The simple leaf morphotypes with the dry mid-Cretaceous regional vegetation are small with peltate blades, and with petioles that recognized by Schrank (1994 and elsewhere) the are either short and twisted (Aryskumia) or long, with Nammoura assemblage is relatively rich in ferns and a distinct abscission line (Nammourophyllum). These conifers suggesting that aridity was not severe, features typically are associated in deciduous angio- perhaps expressed in a fairly dry mid-summer season. sperm leaves. The Nammouran biserrate leaves are 798 V. Krassilov and F. Bacchia among the earliest records of this morphotype that in River Formation of Utah and Colorado. Review of Palaeobotany most regions of the world first appeared no earlier and Palynology 80, 305–310. Chitaley, S. D. & Paradkar, S. A. 1972. Rodeites sahnii reinvesti- than the Turonian (Krassilov, 1997). gated, I. Botanical Journal of Linnaean Society 65, 109–117. It does not seem productive to single out extant Copeland, E. B. 1947. Genera Filicum. The genera of ferns, 247 pp. ecological equivalents because they can be found in (Annales Cryptogamici Phytopathologici). Dal Sasso, C. & Pinna, G. 1997. Aphanizocnemus libanensis various climatic zones. Modern ecological equivalents n. gen. n. sp., a new dolichosaur (Reptilia, Varanoidea) from of the assemblage as a whole may prove more instruc- the Upper Cretaceous of Lebanon. Paleontologia Lombarda 7, tive. In a search for such the following features of the 1–31. Dilcher, D. L. & Basson, P. W. 1990. Mid-Cretaceous angiosperm Nammoura assemblage seem significant: (1) it con- leaves from a new fossil locality in Lebanon. Botanical Gazette tains a subordinate, yet prominent, fern-gymnosperm 151, 538–547. component; (2) the conifer remains, though rare, Dobruskina, I. A. 1997. Turonian plants from the southern Negev, might come from an abundant up-slope source; (3) Israel. Cretaceous Research 18,87–107. El Chair, M., Kerp, H. & Thiedig, F. 1995. Two florules from the compound leaves predominate among angiosperm Jarmah Member of the Early Cretaceous Mesak Formation at morphotypes; (4) the morphology of simple leaves Jabal Tandah south of Awbari and northeast of Sabha, Libya. indicates deciduousness; (5) the leaf blade units (sim- Neues Jahrbuch fu¨r Geologie und Palaöntologie, Monatshafte 1995, 659–670. ple leaves and leaflets of compound leaves) are small, Herendeen, P. S. & Dilcher, D. L. 1990. Fossil mimosoid legumes with xeromorphic features; and (6) the leaf margins from the Eocene and Oligocene of southeastern North America. are serrate in the simple leaves, entire in the leaflets. A Review of Palaeobotany and Palynology 62, 339–361. combination of these features is characteristic of the Hollick, A. 1950. The Upper Cretaceous floras of Alaska. United States Geological Survey, Professional Paper 159,1–116. present day Mediterranean vegetation of the same Huang, Q. C. & Dilcher, D. L. 1994. Evolutionary and paleoeco- territory (Krassilov, 1975, 1984). logical implications of fossil plants from the Lower Cretaceous Sedimentological features of the locality may add to Cheyenne Sandstone of the Western Interior. Geological Society of America, Special Paper 287, 129–144. the climatic interpretation of the floristic assemblage. Hu¨ckel, U. 1970. Die Fischschiefer von Haqel and Hjoula in der The fish beds of Lebanon are considered to be Oberkreide des Libanon. Neues Jahrbuch fu¨r Geologie und Palaön- deposits of an anoxic sedimentary environment, the tologie, Abhandlungen 135, 113–149. Kimura, T. & Huriuchi, J. 1978. Pseudolarix nipponica sp. nov. from origin of which was ascribed to a number of tectonic the Paleogene Noda Group, north-east Japan. Proceedings of the and/or palaeogeographic factors, such as synsedimen- Japan Academy of Sciences 54, 429–434. tary downfaulting or upwelling (recently reviewed in Knobloch, E. 1978. On some primitive angiosperm leaves from the Upper Cretaceous of the Bohemian Massif. Palaeontographica B Schram et al., 1999). An alternative explanation is 166,83–98. prompted by the transported plant material that was Krassilov, V. A. 1975. Paleoecology of terrestrial plants, 230 pp. found not only at Nammoura but also at Haquel from (Wiley, New York). Krassilov, V. A. 1982. Early Cretaceous flora of Mongolia. which indeterminable plant remains were reported by Palaeontographica B 181,1–43. Hu¨ckel (1970). Their constant, though scattered, Krassilov, V. A. 1984. Albian–Cenomanian flora of the Kacha- occurrence indicates a considerable influx of terres- Bodrak intershed, Crimea. Byulleten’ Moskovskogo Obstchestva Ispytateley Prirody, Geologia 59, 104–112. [In Russian] trial organic debris. A dilution of marine surface Krassilov, V. A. 1997. Angiosperm origins: morphological and ecologi- waters by a discharge of fresh waters carrying organic cal aspects, 270 pp. (Pensoft, Sofia). material might have caused anoxia down the water Krassilov, V. A. & Dobruskina, I. A. 1998. A gramminoid plant from the Cretaceous of Middle East. Paleontological Zhurnal column. In the Mediterranean Sea such anoxic events (Moscow) 1998 (4), 106–110. correlate with increased mid-latitude precipitation Krassilov, V. A. & Golovneva, L. B. 1999. A new heterosporous (Rossignol-Strich et al., 1982). At the same time, the plant from the Cretaceous of West Siberia. Review of Palaeobotany and Palynology 105,75–84. thinly laminated limestones of the Nammoura fish bed Labandeira, C. C., Dilcher, D. L., Davis, D. R. & Wagner, D. L. indicate a pronounced seasonality of precipitation, 1994. Ninety-seven million years of angiosperm-insect associ- which concurs with palaeoecological aspect of the ation: paleobiological insights into the meaning of coevolution. Proceedings of the National Academy of Sciences, USA 91, 12,278– plant assemblage. 12,282. Manchester, S. R. 1992. Flowers, fruits, and pollen of Florissantia, an extinct malvalean genus from the Eocene and Oligocene of western North America. American Journal of Botany 79, 996–1008. References Mosley, M. F. 1972. Morphological studies of the Nymphaeaceae. VI. Development of flower of Nuphar. Phytomorphology 21, Barale, G., Philippe, M., Tayech-Mannal, B. & Zarbout, M. 1997. 253–283. Dećouverte d’un flore a`pte´ridophytes et gymnospermes dans le Neˇmejc, F. & Kvacˇek, Z. 1975. Senonian plant macrofossils from Cre´tace´ infe´rieur de la re´gion de Tataouine (Sud tunisien). the region of Zliv and Hluboka´ (near Eske´ Budeˇjovice) in South Comptes Rendus de l’Academie des Sciences, Paris, Sciences de la Bohemia, 82 pp. (Universita Karlova, Praha). Terre et des Plane`tes 326, 221–224. Rossignol-Strich, M., Nesteroff, W. & Olive, P. 1982. After the Call, V. B. & Dilcher, D. L. 1994. Parvileguminophyllum coloraden- deluge: Mediterranean stagnation and sapropel formation. Nature sis, a new combination for Mimosites coloradensis Knowlton, Green 295, 105–110. Cenomanian florule of Nammoura, Lebanon 799

Rothwell, G. W. & Stockey, R. A. 1994. The role of Hydropteris Upchurch, G. R. 1984. Cuticle evolution in Early Cretaceous pinnata gen. et sp. nov. in reconstructing the cladistics of angiosperms from the Potomac Group of Virginia and Maryland. heterosporous ferns. American Journal of Botany 81, 479–492. Annals of the Missouri Botanical Garden 71, 522–550. Saporta, G. 1868. Prodrome d’une flore fossile des travertines Upchurch, G. R. & Dilcher, D. L. 1990. Cenomanian angiosperm anciens de Se´zanne. Me´moire de la Socie´te´ Geólogique de France 28, leaf megafossils, Dakota Formation, Rose Creek Locality, 289–436. Jefferson County, southeastern Nebraska. United States Geological Schram, F. R., Hof, C. H. & Steeman, F. A. 1999. Thylacocephala Survey, Bulletin 1915,1–55. (Arthropoda: Crustaceae?) from the Cretaceous of Lebanon and Watson, J. & Harrison, N. A. 1998. Abietites linkii (Ro¨mer) and implications for thylacocephalan systematics. Palaeontology 42, Pseudotorellia heterophylla Watson: coniferous or ginkgoalean? 769–799. Cretaceous Research 19, 239–278. Schrank, E. 1994. Nonmarine Cretaceous palynology of northern Wing, S. L. & Hickey, L. J. 1984. The Platycarya perplex and the Kordofan, Sudan, with notes on fossil Salviniales (water ferns). evolution of the Juglandaceae. American Journal of Botany 71, Geologische Rundschau 83, 773–786. 388–411. Shilin, P. V. 1986. Late Cretaceous floras of Kazakhstan: systematics, geological history and stratigraphic significance, 200 pp. (Nauka, Alma-Ata).