- Email: [email protected]
Brachyphyllum lorchii sp. nov. from the upper jurassic of Israel
Review of Palaeobotany and Palynology, 46 (1986): 227--234 Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands
BRACHYPHYLLUM OF ISRAEL
227
L O R C H I I SP. N O V . F R O M T H E U P P E R J U R A S S I C
MENAHEM RAAB', AHARON HOROWITZ 2 and BRIAN H. CONWAY' 1Geological Survey of Israel, 30 Malkhe Israel Str., Jerusalem (Israel) 2Departrnent of Geophysics and Planetary Sciences, Tel Aviv University, Tel Aviv (Israel) (Received January 2, 1985; revised and accepted May 21, 1985)
ABSTRACT Raab, M., Horowitz, A. and Conway, B.H., 1986. Brachyphyllum lorchii sp. nov. from the Upper Jurassic of Israel. Rev. Palaeobot. Palynol., 46: 227--234. Coniferacean foliage referred to Brachyphyllum lorchii sp. nov. is described from the Jurassic Kidod Formation, recovered from core No.1 of Kidod 2 well, drilled W of the Dead Sea, Israel. The new species is differentiated from other species of the genus Brachyphyllum mainly by the shape of its leaves. Dispersed pollen grains referable to Araucariacites australis Cookson and Callialasporites trilobatus (Balme), found in intimate association with the shoots, are thought to have been produced by this plant. INTRODUCTION Several o c c u r r e n c e s o f B r a c h y p h y l l u m are r e p o r t e d b y L o r c h ( 1 9 6 3 ; 1 9 6 7 a , b; 1 9 6 8 ) f r o m t h e Jurassic r o c k s c r o p p i n g o u t in anticlinal erosion cirques in s o u t h e r n Israel a n d in Sinai. O f these, t h e m o s t p r o d u c t i v e is t h e E a r l y Jurassic M i s h h o r F o r m a t i o n in M a k h t e s h R a m o n , f r o m w h i c h L o r c h {1968) h a d i d e n t i f i e d several n e w species o f B r a c h y p h y l l u m : B. negevensis, B. p u l c h e r a n d B. p o r r i g e n t e . T h e Middle Jurassic o f Sinai c o n t a i n e d B. o b e s u m a n d B. sp. ( L o r c h , 196 7a). T h e Middle Jurassic o f M a k h t e s h R a m o n c o n t a i n e d , a m o n g a varied b e n n e t t i t a l e a n a n d ferns flora, B. cf. mamillare ( L o r c h , 1 9 6 7 b ) . B r a c h y p h y l l u m o b e s u m is also r e p o r t e d b y L o r c h ( 1 9 6 3 ) f r o m E a r l y C r e t a c e o u s r o c k s o f M a k h t e s h R a m o n . T h e c o n i f e r a l e a n fossils p r e d o m i n a t e in t h e E a r l y Jurassic o u t c r o p , while all o t h e r a s s o c i a t i o n s are c h a r a c t e r i z e d b y a w e a l t h o f p t e r i d o p h y t e flora, a c c o m p a n i e d b y B e n n e t t i t a l e s . The specimens described here have been recovered from core No.1 of K i d o d 2 b o r e h o l e , drilled in search o f oil in t h e J u d e a n D e s e r t , W o f t h e D e a d S e a ( F i g . l ) . T h e y o c c u r in black, p y r i t i c shales o f t h e K i d o d F o r m a t i o n . Pieces o f t h e b l a c k shale h a v e b e e n r e m o v e d f o r a b e t t e r e x a m i n a t i o n o f t h e s p e c i m e n s , a n d w e r e c o n s e q u e n t l y m a c e r a t e d in h y d r o f l u o r i c acid f o r t h e i r organic c o n t e n t s , cuticles a n d s p o r o m o r p h s , t h a t , t o g e t h e r w i t h t h e fossil p l a n t s , are t h e s u b j e c t o f t h e p r e s e n t s t u d y . 0034-6667/86/$03.50
© 1986 Elsevier Science Publishers B.V.
228
q3¢
5{
i. AV,V
:Ho.
o
,~.
14:
A
D]
E.
mo, SINAI 51] km I
AI
6ULF
OF
ELA]
I
Fig.1. L o c a t i o n m a p . STRATIGRAPHY
Core No.1 of Kidod 2 borehole was taken at the base of the Kidod Formation, at a depth of 1200 m. The core contains abundant megafossils, mainly ammonites and a few pelecypods as well as abundant palynomorphs. It consists of black to dark grey clays and shaley clays, rich in organic matter. The ammonites are small (many of them are nuclei), partly pyritic and badly preserved. However, they show a close resemblance to some of the forms described by Haas (1955) from the Kidod Formation of Mt. Hermon in northern Israel. Such forms as Phylloceras plicatum Neumayr, Campylites freboldi Haas, Sublunuloceras guthei (Noetling), S. socium Haas, Peltoceratoides cf. ardeunense (d'Orbigny) and ?Euaspidoceras rotundatum Haas, could tentatively be identified. The age assigned to the Mt. Hermon assemblage by Haas (Op.cit., p.204) is earliest Oxfordian, suggesting thus a certain Late Callovian to Early Oxfordian age for the Kidod 2 bore hole assemblage. This age was also proposed by Lewy (1983, p.14) for the Kidod Formation in Israel. The size range and the pyritization of the ammonites indicate a rather deep-neritic, quiet and euxinic environment o f deposition (Haas, 1955, p.97; Lewy, 1983). The palynomorphs comprise the following taxa (among others): Ctenidodinium ornatum (Eisenack) Deflandre 1938, Gonyaulacysta jurassica (Deflandre) Norris and Sarjeant 1965, Polystephanephorus calathus (Sarjeant)
229 Downie and Sarjeant 1965 and Polystephanephorus paracalathus (Sarjeant) Downie and Sarjeant 1965, which confirms the Late Callovian to Early Oxfordian age of the Kidod Formation (Sarjeant, 1979). The formation is k n o w n from outcrops and many boreholes over the entire region, from M o u n t Hermon in the north down to Sinai in the south (Goldberg, 1970). It comprises 30--150 m of black shales, in places rich in ammonite fauna. The Kidod Formation overlies the Arad Group of Early and Middle Jurassic age and is overlain by carbonate rocks of the Be'er Sheva Formation, of Late Jurassic (Oxfordian) age. THE PLANT MATERIAL Family ARAUCARIACEAE
Genus Brachyphyllum Lindley and Hutton ex Brongniart 1828 Brachyphyllum Brongniart, p.109 (definition, no type species). 1836 Brachyphyllum Lindley and Hutton, plate 188. 1979 Brachyphyllum Harris, p.4 (emended diagnosis).
Brachyphyllum lorchii sp. nov. (Plate I, 1--6; Plate II, 1--3) Holotype: Type No.1471; Sample No.M-6204; Div. of Paleontology, Geological Survey of Israel. Type locality: Kidod 2 borehole, core N o . l , box 17, 1200 m. Age: Late Callovian to Early Oxfordian. Formation: Kidod Formation. Derivation of name: In h o n o u r of Prof. Yaacov Lorch, The Hebrew University o f Jerusalem, who contributed significantly to palaeobotanical studies in Israel. Description: Leafy shoots -- several specimens recovered from core N o . l , Kidod 2 borehole (Plate I, 1--5), of which one is exceptionally well preserved and serves as the holotype. H o l o t y p e 25 mm long, some 5 m m in diameter, branched at its top end, probably in a pinnate manner. Leaves are arranged helicoidally. Leaves -- attached to the stem by rhomboidal basal cushions 3--3.5 m m wide. The free part of the leaves rises from the entire basal cushion at an angle of some 35 ° outwards and upwards and is 2--3 m m long. The length o f the entire leaf is 4--4.5 mm. The shape of the leaf is rhomboidal, the free part slightly more elongated than the cushion. Leaf apex rounded, typically aquiliform (Plate I, 6; Plate II, 1, 2), sides approaching apex at an angle of some 80 °. Leaf surface distinctly furrowed by stomatal grooves, clearly seen on the entire adaxial surface, leaf margins entire. The central adaxial part of the leaf bears a prominent ridge t h a t curves inwards towards the apex. Cuticle -- the coarse structure of the cuticle and stomata can be seen in the scanning electron micrographs of the h o l o t y p e (Plate II, 3), but are
230
PLATE
4
I
5
231
much clearer in slides prepared for light microscope examination from pieces of shale removed for cleaning the specimen (Plate II, 4). It seems though that most cuticle pieces obtained in this way are from the adaxial face, while the abaxial is probably not represented, the material was macerated in hydrofluoric acid, washed in water and m o u n t e d in synthetic canada balsam. This treatment did n o t cause distortions but the cuticular material seems to have expanded some 40--50%, as was observed while measuring the stomata with the SEM and comparing with the photomicrographs. The measurements given here are corrected, but the scale in Plate II is the same as observed through the microscope. The cuticle is some 4--5 g m thick, epidermal cells typically equidimensional, some 20 p m across, mostly forming longitudinal rows. Cell walls are quite broad, corners of cells mostly rounded. Surface of cells smooth, occasionally very faintly granulate. Stomataequally c o m m o n on all specimens observed, but as stated above it seems that the material only represents adaxial surfaces. Stomata arranged in rows occupying stomatal grooves, which give the leaves a typical striated appearance. S t o m a t a mostly circular, sometimes slightly elongated parallel to the leaf axis, mostly some 20 p m across. Guard cells rarely preserved, typically encircled by four subsidiary cells ridged at the inner and o u t e r circumferences. The entire stomatal apparatus is slightly sunken in the epidermis, forming a wide pit, mostly equidimensional, some 40 pm across, rarely somewhat longitudinally elongated, up to 50 X 30 #m. The encircling subsidiary cells unspecialized, with no papillae, the polar and lateral ones mostly similar. Discussion: the typical aquiline form of the leaf, with its prominent ridge and striation are different from those of other species of Brachyphyllum described by Kendall (1947, 1949), Lorch (1967a, b, 1968) and Harris (1979). This justifies the erection of a new species, namely B. lorchii. It seems almost b e y o n d d o u b t that it belongs, as other species of the same genus (Stockey, 1982), to the Araucariaceae, especially from its cuticle structure. THE P O L L E N I F E R O U S M A T E R I A L
The chips removed from the piece of black shale containing B. lorchii have been treated in the same manner for obtaining palynomorphs as described above for the cuticles, b u t only the finer fraction was used to prepare the slides. We do n o t have any means of control on possible size changes due to preparation, since no palynomorphs have been observed in the SEM, therefore no correction is given for the measurements. It is of course impossible to infer from the behavior of cuticles that of sporopollenin. PLATE
I
1--6. Brachyphyllum lorchii sp. nov. 1--2; Holotype; 1, A m m o n i u m chloride-coated; 2, Gold-coated; 3--5, Other specimens from the same horizon. Scale for 1--5: 5 ram; 6, Detail o f the holotype; scale 1 mm., SEM by M. Dvorachek.
232
PLATE II
I
im
J
233 Only two ty pe s o f pollen (Plate II, 5, 6) com pose t he majority (m ore than 80%) o f the spectrum obt ai ned f r om t h e plant sample - - A r a u c a r i a c i t e s australis which dominates, and Callialasporites trilobatus which composes some 10% o f the spectrum. GYMNOSPERMAE Family ARAUCARIACEAE Genus Araucariacites C ooks on ex Couper 1947 Araucariacites Cookson, p.130, plate 13, figs.l--4. 1953 Araucariacites Couper, p.39. Araucariacites australis Cookson, 1947 Spherical, inaperturate pollen grain. Intectate, psilate to sub-granulate. Exine fine -- a b o u t 1 /~m thick. Grain occasionally folded and fractured. Size: a p p r o x i m a t e l y 80 p m across (10 specimens). It should be n o t e d t hat in some o th er Jurassic samples (Horowitz, 1970) the average size was only 50 ~m, so it might well be t hat t he grains also swelled during maceration. Genus Callialasporites Sukh Dev 1961 CaUialasporites Sukh Dev, p.48. Callialasporites trilobatus (Balme) Sukh Dev, 1961 Cavate, inaperturate pollen grain. B o d y central, subtriangular, exine a b o u t 2 ~m thick, finely granulated. Trilobate equatorial bladder attached to b od y , 6--10 ~m wide, finely granulated, sometimes with radial folds. Size: 60 ~m (body) ; 7 0 - - 7 5 p m (overall) (5 specimens). DISCUSSION The intimate association of the B. lorchii shoots and the pollen Araucariacites australis and Callialasporites trilobatus seem to strengthen opinions by Kendall (1949), Harris (1979) and S t o c k e y (1982) as to the natural relations PLATE II 1--3. Details of the leaves, holotype, SEM by M. Dvorachek. 1, 2, Single leaves, showing the typical aquiliform shapes, the striations and the arrangement of stomata. Scale: 0.5 ram. 3, Stomata, scale 10 am. 4. Cuticle under a transmitted light microscope, showing a row of stomata, in one of them the guard cells are preserved. 5. Araucariacites australis under a light microscope. 6. Callialasporites trilobatus under a light microscope. Scale for 4--6:10 ~m.
234 of those pollen grains and the fossil Araucariaceae, although unfortunately n o m a l e c o n e s h a v e b e e n f o u n d in o u r s a m p l e s . T h e c o n n e c t i o n b e t w e e n Brachyphyllum a n d Araucariacites--Callialasporites ( V a n K o n i j n e n b u r g - V a n C i t t e r t , 1 9 7 1 ) is f u r t h e r s u p p o r t e d b y t h e r e s u l t s o f t h e p r e s e n t s ~ u d y . ACKNOWLEDGEMENTS T h e a u t h o r s w i s h t o t h a n k Mr. M. D v o r a c h e k f o r t a k i n g t h e S E M p h o t o graphs. REFERENCES Goldberg, M., 1970. The lithostratigraphy of Arad group (Jurassic) in the northern Negev. Geol. Surv. Isr., Jerusalem, Rep., MM/3/70, 137 pp. (In Hebrew, English summary). Haas, O., 1955. Revision of the Jurassic ammonite fauna of Mount Hermon, Syria. Bull. Am. Mus. Nat. Hist., 108: 210. Harris, T.M., 1979. The Yorkshire Jurassic flora, V. Coniferales. Br. Mus. (Nat. Hist.) Publ., 83, 166 pp. Horowitz, A., 1970. Jurassic micro flora from the northern Negev, Israel. Isr. J. Earth Sci., 19: 153--182. Kendall, M.W., 1947. On five species of Brachyphyllum from the Jurassic of Yorkshire and Wiltshire. Ann. Mag. Nat. Hist. Ser. 2, XIV: 255. Kendall, M.W., 1949. A Jurassic member of the Araucariaceae. Ann. Bot. N.S. XIII, 50: 151--161.
Lewy, Z., 1983. Upper Callovian ammonites and Middle Jurassic geological history of the Middle East. Isr. Geol. Surv. Bull., 76 : 16--56. Lorch, J., 1963. The fossil flora of Makhtesh Ramon. Proc. Isr. Geol. Soc., 15. Lorch, J., 1967a. A Jurassic florule from Sinai. Isr. J. Bot., 16: 29--37. Lorch, J., 1967b. A Jurassic flora of Makhtesh Ramon, Israel. Isr. J. Bot., 16: 131--155. Lorch, J., 1968. Some Jurassic conifers from Israel. J. Linn. Soc. London. Bot., 61: 177--188. Sarjeant, W.A.S., 1979. Middle and Upper Jurassic dinoflagellate cysts: The world excluding North America. Am. Assoc. Stratigr. Palynol., Contrib. Ser., 5B: 133--151. Stockey, R.A., 1982. The Araucariaceae: an evolutionary perspective. Rev. Palaeobot. Palynol., 37: 133--154. Sukh Dev, 1961. The fossil flora of the Jabalpur Series 3. Spores and pollen grains. Palaeobotanist, 8 : 43--56. Van Konijnenburg - - Van Cittert, J.H.A., 1971. In situ gymnosperm pollen from the middle Jurassic of Yorkshire. Acta Bot. Neerl., 20: 1--95.
BRACHYPHYLLUM OF ISRAEL
227
L O R C H I I SP. N O V . F R O M T H E U P P E R J U R A S S I C
MENAHEM RAAB', AHARON HOROWITZ 2 and BRIAN H. CONWAY' 1Geological Survey of Israel, 30 Malkhe Israel Str., Jerusalem (Israel) 2Departrnent of Geophysics and Planetary Sciences, Tel Aviv University, Tel Aviv (Israel) (Received January 2, 1985; revised and accepted May 21, 1985)
ABSTRACT Raab, M., Horowitz, A. and Conway, B.H., 1986. Brachyphyllum lorchii sp. nov. from the Upper Jurassic of Israel. Rev. Palaeobot. Palynol., 46: 227--234. Coniferacean foliage referred to Brachyphyllum lorchii sp. nov. is described from the Jurassic Kidod Formation, recovered from core No.1 of Kidod 2 well, drilled W of the Dead Sea, Israel. The new species is differentiated from other species of the genus Brachyphyllum mainly by the shape of its leaves. Dispersed pollen grains referable to Araucariacites australis Cookson and Callialasporites trilobatus (Balme), found in intimate association with the shoots, are thought to have been produced by this plant. INTRODUCTION Several o c c u r r e n c e s o f B r a c h y p h y l l u m are r e p o r t e d b y L o r c h ( 1 9 6 3 ; 1 9 6 7 a , b; 1 9 6 8 ) f r o m t h e Jurassic r o c k s c r o p p i n g o u t in anticlinal erosion cirques in s o u t h e r n Israel a n d in Sinai. O f these, t h e m o s t p r o d u c t i v e is t h e E a r l y Jurassic M i s h h o r F o r m a t i o n in M a k h t e s h R a m o n , f r o m w h i c h L o r c h {1968) h a d i d e n t i f i e d several n e w species o f B r a c h y p h y l l u m : B. negevensis, B. p u l c h e r a n d B. p o r r i g e n t e . T h e Middle Jurassic o f Sinai c o n t a i n e d B. o b e s u m a n d B. sp. ( L o r c h , 196 7a). T h e Middle Jurassic o f M a k h t e s h R a m o n c o n t a i n e d , a m o n g a varied b e n n e t t i t a l e a n a n d ferns flora, B. cf. mamillare ( L o r c h , 1 9 6 7 b ) . B r a c h y p h y l l u m o b e s u m is also r e p o r t e d b y L o r c h ( 1 9 6 3 ) f r o m E a r l y C r e t a c e o u s r o c k s o f M a k h t e s h R a m o n . T h e c o n i f e r a l e a n fossils p r e d o m i n a t e in t h e E a r l y Jurassic o u t c r o p , while all o t h e r a s s o c i a t i o n s are c h a r a c t e r i z e d b y a w e a l t h o f p t e r i d o p h y t e flora, a c c o m p a n i e d b y B e n n e t t i t a l e s . The specimens described here have been recovered from core No.1 of K i d o d 2 b o r e h o l e , drilled in search o f oil in t h e J u d e a n D e s e r t , W o f t h e D e a d S e a ( F i g . l ) . T h e y o c c u r in black, p y r i t i c shales o f t h e K i d o d F o r m a t i o n . Pieces o f t h e b l a c k shale h a v e b e e n r e m o v e d f o r a b e t t e r e x a m i n a t i o n o f t h e s p e c i m e n s , a n d w e r e c o n s e q u e n t l y m a c e r a t e d in h y d r o f l u o r i c acid f o r t h e i r organic c o n t e n t s , cuticles a n d s p o r o m o r p h s , t h a t , t o g e t h e r w i t h t h e fossil p l a n t s , are t h e s u b j e c t o f t h e p r e s e n t s t u d y . 0034-6667/86/$03.50
© 1986 Elsevier Science Publishers B.V.
228
q3¢
5{
i. AV,V
:Ho.
o
,~.
14:
A
D]
E.
mo, SINAI 51] km I
AI
6ULF
OF
ELA]
I
Fig.1. L o c a t i o n m a p . STRATIGRAPHY
Core No.1 of Kidod 2 borehole was taken at the base of the Kidod Formation, at a depth of 1200 m. The core contains abundant megafossils, mainly ammonites and a few pelecypods as well as abundant palynomorphs. It consists of black to dark grey clays and shaley clays, rich in organic matter. The ammonites are small (many of them are nuclei), partly pyritic and badly preserved. However, they show a close resemblance to some of the forms described by Haas (1955) from the Kidod Formation of Mt. Hermon in northern Israel. Such forms as Phylloceras plicatum Neumayr, Campylites freboldi Haas, Sublunuloceras guthei (Noetling), S. socium Haas, Peltoceratoides cf. ardeunense (d'Orbigny) and ?Euaspidoceras rotundatum Haas, could tentatively be identified. The age assigned to the Mt. Hermon assemblage by Haas (Op.cit., p.204) is earliest Oxfordian, suggesting thus a certain Late Callovian to Early Oxfordian age for the Kidod 2 bore hole assemblage. This age was also proposed by Lewy (1983, p.14) for the Kidod Formation in Israel. The size range and the pyritization of the ammonites indicate a rather deep-neritic, quiet and euxinic environment o f deposition (Haas, 1955, p.97; Lewy, 1983). The palynomorphs comprise the following taxa (among others): Ctenidodinium ornatum (Eisenack) Deflandre 1938, Gonyaulacysta jurassica (Deflandre) Norris and Sarjeant 1965, Polystephanephorus calathus (Sarjeant)
229 Downie and Sarjeant 1965 and Polystephanephorus paracalathus (Sarjeant) Downie and Sarjeant 1965, which confirms the Late Callovian to Early Oxfordian age of the Kidod Formation (Sarjeant, 1979). The formation is k n o w n from outcrops and many boreholes over the entire region, from M o u n t Hermon in the north down to Sinai in the south (Goldberg, 1970). It comprises 30--150 m of black shales, in places rich in ammonite fauna. The Kidod Formation overlies the Arad Group of Early and Middle Jurassic age and is overlain by carbonate rocks of the Be'er Sheva Formation, of Late Jurassic (Oxfordian) age. THE PLANT MATERIAL Family ARAUCARIACEAE
Genus Brachyphyllum Lindley and Hutton ex Brongniart 1828 Brachyphyllum Brongniart, p.109 (definition, no type species). 1836 Brachyphyllum Lindley and Hutton, plate 188. 1979 Brachyphyllum Harris, p.4 (emended diagnosis).
Brachyphyllum lorchii sp. nov. (Plate I, 1--6; Plate II, 1--3) Holotype: Type No.1471; Sample No.M-6204; Div. of Paleontology, Geological Survey of Israel. Type locality: Kidod 2 borehole, core N o . l , box 17, 1200 m. Age: Late Callovian to Early Oxfordian. Formation: Kidod Formation. Derivation of name: In h o n o u r of Prof. Yaacov Lorch, The Hebrew University o f Jerusalem, who contributed significantly to palaeobotanical studies in Israel. Description: Leafy shoots -- several specimens recovered from core N o . l , Kidod 2 borehole (Plate I, 1--5), of which one is exceptionally well preserved and serves as the holotype. H o l o t y p e 25 mm long, some 5 m m in diameter, branched at its top end, probably in a pinnate manner. Leaves are arranged helicoidally. Leaves -- attached to the stem by rhomboidal basal cushions 3--3.5 m m wide. The free part of the leaves rises from the entire basal cushion at an angle of some 35 ° outwards and upwards and is 2--3 m m long. The length o f the entire leaf is 4--4.5 mm. The shape of the leaf is rhomboidal, the free part slightly more elongated than the cushion. Leaf apex rounded, typically aquiliform (Plate I, 6; Plate II, 1, 2), sides approaching apex at an angle of some 80 °. Leaf surface distinctly furrowed by stomatal grooves, clearly seen on the entire adaxial surface, leaf margins entire. The central adaxial part of the leaf bears a prominent ridge t h a t curves inwards towards the apex. Cuticle -- the coarse structure of the cuticle and stomata can be seen in the scanning electron micrographs of the h o l o t y p e (Plate II, 3), but are
230
PLATE
4
I
5
231
much clearer in slides prepared for light microscope examination from pieces of shale removed for cleaning the specimen (Plate II, 4). It seems though that most cuticle pieces obtained in this way are from the adaxial face, while the abaxial is probably not represented, the material was macerated in hydrofluoric acid, washed in water and m o u n t e d in synthetic canada balsam. This treatment did n o t cause distortions but the cuticular material seems to have expanded some 40--50%, as was observed while measuring the stomata with the SEM and comparing with the photomicrographs. The measurements given here are corrected, but the scale in Plate II is the same as observed through the microscope. The cuticle is some 4--5 g m thick, epidermal cells typically equidimensional, some 20 p m across, mostly forming longitudinal rows. Cell walls are quite broad, corners of cells mostly rounded. Surface of cells smooth, occasionally very faintly granulate. Stomataequally c o m m o n on all specimens observed, but as stated above it seems that the material only represents adaxial surfaces. Stomata arranged in rows occupying stomatal grooves, which give the leaves a typical striated appearance. S t o m a t a mostly circular, sometimes slightly elongated parallel to the leaf axis, mostly some 20 p m across. Guard cells rarely preserved, typically encircled by four subsidiary cells ridged at the inner and o u t e r circumferences. The entire stomatal apparatus is slightly sunken in the epidermis, forming a wide pit, mostly equidimensional, some 40 pm across, rarely somewhat longitudinally elongated, up to 50 X 30 #m. The encircling subsidiary cells unspecialized, with no papillae, the polar and lateral ones mostly similar. Discussion: the typical aquiline form of the leaf, with its prominent ridge and striation are different from those of other species of Brachyphyllum described by Kendall (1947, 1949), Lorch (1967a, b, 1968) and Harris (1979). This justifies the erection of a new species, namely B. lorchii. It seems almost b e y o n d d o u b t that it belongs, as other species of the same genus (Stockey, 1982), to the Araucariaceae, especially from its cuticle structure. THE P O L L E N I F E R O U S M A T E R I A L
The chips removed from the piece of black shale containing B. lorchii have been treated in the same manner for obtaining palynomorphs as described above for the cuticles, b u t only the finer fraction was used to prepare the slides. We do n o t have any means of control on possible size changes due to preparation, since no palynomorphs have been observed in the SEM, therefore no correction is given for the measurements. It is of course impossible to infer from the behavior of cuticles that of sporopollenin. PLATE
I
1--6. Brachyphyllum lorchii sp. nov. 1--2; Holotype; 1, A m m o n i u m chloride-coated; 2, Gold-coated; 3--5, Other specimens from the same horizon. Scale for 1--5: 5 ram; 6, Detail o f the holotype; scale 1 mm., SEM by M. Dvorachek.
232
PLATE II
I
im
J
233 Only two ty pe s o f pollen (Plate II, 5, 6) com pose t he majority (m ore than 80%) o f the spectrum obt ai ned f r om t h e plant sample - - A r a u c a r i a c i t e s australis which dominates, and Callialasporites trilobatus which composes some 10% o f the spectrum. GYMNOSPERMAE Family ARAUCARIACEAE Genus Araucariacites C ooks on ex Couper 1947 Araucariacites Cookson, p.130, plate 13, figs.l--4. 1953 Araucariacites Couper, p.39. Araucariacites australis Cookson, 1947 Spherical, inaperturate pollen grain. Intectate, psilate to sub-granulate. Exine fine -- a b o u t 1 /~m thick. Grain occasionally folded and fractured. Size: a p p r o x i m a t e l y 80 p m across (10 specimens). It should be n o t e d t hat in some o th er Jurassic samples (Horowitz, 1970) the average size was only 50 ~m, so it might well be t hat t he grains also swelled during maceration. Genus Callialasporites Sukh Dev 1961 CaUialasporites Sukh Dev, p.48. Callialasporites trilobatus (Balme) Sukh Dev, 1961 Cavate, inaperturate pollen grain. B o d y central, subtriangular, exine a b o u t 2 ~m thick, finely granulated. Trilobate equatorial bladder attached to b od y , 6--10 ~m wide, finely granulated, sometimes with radial folds. Size: 60 ~m (body) ; 7 0 - - 7 5 p m (overall) (5 specimens). DISCUSSION The intimate association of the B. lorchii shoots and the pollen Araucariacites australis and Callialasporites trilobatus seem to strengthen opinions by Kendall (1949), Harris (1979) and S t o c k e y (1982) as to the natural relations PLATE II 1--3. Details of the leaves, holotype, SEM by M. Dvorachek. 1, 2, Single leaves, showing the typical aquiliform shapes, the striations and the arrangement of stomata. Scale: 0.5 ram. 3, Stomata, scale 10 am. 4. Cuticle under a transmitted light microscope, showing a row of stomata, in one of them the guard cells are preserved. 5. Araucariacites australis under a light microscope. 6. Callialasporites trilobatus under a light microscope. Scale for 4--6:10 ~m.
234 of those pollen grains and the fossil Araucariaceae, although unfortunately n o m a l e c o n e s h a v e b e e n f o u n d in o u r s a m p l e s . T h e c o n n e c t i o n b e t w e e n Brachyphyllum a n d Araucariacites--Callialasporites ( V a n K o n i j n e n b u r g - V a n C i t t e r t , 1 9 7 1 ) is f u r t h e r s u p p o r t e d b y t h e r e s u l t s o f t h e p r e s e n t s ~ u d y . ACKNOWLEDGEMENTS T h e a u t h o r s w i s h t o t h a n k Mr. M. D v o r a c h e k f o r t a k i n g t h e S E M p h o t o graphs. REFERENCES Goldberg, M., 1970. The lithostratigraphy of Arad group (Jurassic) in the northern Negev. Geol. Surv. Isr., Jerusalem, Rep., MM/3/70, 137 pp. (In Hebrew, English summary). Haas, O., 1955. Revision of the Jurassic ammonite fauna of Mount Hermon, Syria. Bull. Am. Mus. Nat. Hist., 108: 210. Harris, T.M., 1979. The Yorkshire Jurassic flora, V. Coniferales. Br. Mus. (Nat. Hist.) Publ., 83, 166 pp. Horowitz, A., 1970. Jurassic micro flora from the northern Negev, Israel. Isr. J. Earth Sci., 19: 153--182. Kendall, M.W., 1947. On five species of Brachyphyllum from the Jurassic of Yorkshire and Wiltshire. Ann. Mag. Nat. Hist. Ser. 2, XIV: 255. Kendall, M.W., 1949. A Jurassic member of the Araucariaceae. Ann. Bot. N.S. XIII, 50: 151--161.
Lewy, Z., 1983. Upper Callovian ammonites and Middle Jurassic geological history of the Middle East. Isr. Geol. Surv. Bull., 76 : 16--56. Lorch, J., 1963. The fossil flora of Makhtesh Ramon. Proc. Isr. Geol. Soc., 15. Lorch, J., 1967a. A Jurassic florule from Sinai. Isr. J. Bot., 16: 29--37. Lorch, J., 1967b. A Jurassic flora of Makhtesh Ramon, Israel. Isr. J. Bot., 16: 131--155. Lorch, J., 1968. Some Jurassic conifers from Israel. J. Linn. Soc. London. Bot., 61: 177--188. Sarjeant, W.A.S., 1979. Middle and Upper Jurassic dinoflagellate cysts: The world excluding North America. Am. Assoc. Stratigr. Palynol., Contrib. Ser., 5B: 133--151. Stockey, R.A., 1982. The Araucariaceae: an evolutionary perspective. Rev. Palaeobot. Palynol., 37: 133--154. Sukh Dev, 1961. The fossil flora of the Jabalpur Series 3. Spores and pollen grains. Palaeobotanist, 8 : 43--56. Van Konijnenburg - - Van Cittert, J.H.A., 1971. In situ gymnosperm pollen from the middle Jurassic of Yorkshire. Acta Bot. Neerl., 20: 1--95.