Novitate MUSEUM vitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. Number 2908, pp. 1-12, figs. 1-6, tables 1-3 10024 February 11, 1988 Goliathyris lewyi, New Species (Brachiopoda, Terebratulacea) from the Jurassic of Gebel El-Minshera, Northern Sinai HOWARD R. FELDMAN! AND ELLIS F. OWEN? ABSTRACT Goliathyris lewyi, new genus and species is de- scribed from the Lamberticeras lamberti Zone, Ju- rassic (Upper Callovian) of Gebel El-Minshera, northern Sinai. Homeomorphic with Aulacothyris spp., G. lewyi is questionably assigned to the fam- ily Dyscoliidae based on its internal resemblance to Trigonithyris, but resembles a zeilleriid exter- nally. INTRODUCTION The present study is part of a preliminary investigation of the brachiopod faunas of the northern Sinai undertaken by us and col- leagues in the Geological Survey of Israel. Our long-range goal is to complete a taxo- nomic revision of the brachiopod faunas of northern Sinai as well as those of Arabia which will help us establish the early history of bra- chiopod species and their evolution within the “Ethiopian” Faunal Province. Analysis of present data supports our contention that this province was invaded by brachiopods migrating from the north in Early Jurassic times which were isolated for the remainder of the Jurassic. These faunas are thought to have subsequently developed special mor- phological characteristics which distinguish them from their original stock. In addition, we are investigating the distribution of bra- chiopod species across faunal realm and province boundaries, specifically the Indo- African Faunal Realm which is now widely dispersed on various continental fragments. The Sinai is situated at the northern part of ' Research Associate, Department of Invertebrates, American Museum of Natural History; Associate Professor, Biology Department, Touro College, New York, New York 10036. ? 10 The Causeway, Horsham, Sussex, England. Copyright © American Museum of Natural History 1988 ISSN 0003-0082 / Price $2.00 2 AMERICAN MUSEUM NOVITATES NO. 2908 SINAI GEBEL EL-MINSHERA METERS 15 -" LOWER CRETACEOUS. "SANDSTONES .”".” O YY ALLOVIAN NSE LIMESTONE: SEE GIOOLASTI LIMESTONE FI ae as ne oe . . ee . ° oe «oe Ke ee He . : . <- A ee . . . . se . . eo. . . cy aoe tee . se « <2 e . 8 ew Oe Se er te i eee ok ry . s y . . lg . . - 7 e ee ° ce ee q a . . . te 7 . . . oe ° . e ih aes . . . ry a oe . eee . . e - . ° . s % oe 4 : . . . * id a . . = . : . . . 4 . . . é . Poe i 5 SLIMY SANDSTONE] 227 0. oe Fig. 1. Locality map of northern Sinai (inset) and detailed lithologic section of outcrop area. The triangle represents approximate strata from which Goliathyris lewyi, new species, was collected. this realm and consequently the brachiopods Taxonomic revision of the Sinai brachio- are very likely to include species belonging pods will also enable us to define, with greater to the equatorial Tethyan Realm. accuracy, faunal realm and province bound- 1988 FELDMAN AND OWEN: GOLIATHYRIS LEWYI ioe) TABLE 1 Stratigraphic Distribution of Some Jurassic Ammonites at Gebel El-Minshera, Northern Sinai, and Correlation with Ammonite Faunas in Central Saudi Arabia Unnamed stratigraphic units, Gebel El-Minshera Cretaceous Sandstone —_ UNGONFORMITY <-2-02-62-tesete-cepee--epeeeety ites Jurassic (European Upper Callovian) Gray-black dense limestone Marl and clay “*Clydoniceras” (with G. lewyi) Quenstedtoceras Pachyerymnoceras Bioclastic limestone Paracenoceras Limy sandstone — aries. The “‘Ethiopian”’ Faunal Province, for example, is recognizable from early in the Jurassic until the middle and possibly the end of the Cretaceous by the presence of endemic taxa at the species, genus, and family level. These endemics have been recognized in the ammonoid cephalopoda (Arkell, 1952, 1956), in the trigoniacean and crassatellacean bi- valves (Kitchin, 1912), and particularly in the brachiopods (Weir, 1925; Muir-Wood, 1935). This same province has been recognized in India, East Africa, and Madagascar and at the end of the Jurassic in South America; it may also extend eastward as far as New Cal- edonia. Its first occurrence seems to be in the shallow seas following rifts formed during the breakup of Gondwanaland but is apparently limited at an unknown southern margin, as none of its species are known in the geosyn- clinal contemporaneous deposits of Antarc- tica or New Zealand. During much of the Early Jurassic, ubiquitous genera such as Tet- rarhynchia, Lobothyris, and Zeilleria, found in the faunas of most known outcrops, be- came so generalized that it is difficult to plot them in terms of distribution (Ager, 1973). The description and revision of specialized forms in particular will facilitate the delinea- tion of province boundaries as well as the recognition of faunal realms and distribution of genera. The Goliathyris described herein as G. lew- yi, new genus and species, was collected from Ammonites (this report) Ammonite faunas Ammonites (Imlay, 1970) (Arkell, 1952) Lamberticeras lamberti Zone Peltoceras athleta Zone Pachyerymnoceras Pachyceras Erymnoceras Erymnoceras loose debris on a slope of marl and clay (fig. 1) of Upper Callovian age along with Pachy- erymnoceras, Quenstedtoceras, and ““Clydon- iceras”’ pseudodiscus Arkell (Cephalopoda). The marly unit is overlain by gray-black dense limestone, also of Upper Callovian age, con- taining within its matrix specimens of Pel- toceras trifidum (Quenstedt), ‘‘Clydonice- ras,”’ and Pachyerymnoceras (Cephalopoda); Putealiceras, Pseudomelania (Gastropoda), and unidentified corals. The top of the unit lies unconformably under a series of Lower Cretaceous sandstones. A thin, bioclastic limestone unit containing Pachyerymnoceras and Paracenoceras prohexagonum (Cepha- lopoda) directly underlies the marl and clay unit from which the Goliathyris specimens were collected. The age of the upper part of the Jurassic strata at Gebel El-Minshera appears to be Upper Callovian (table 1) on the basis of am- monites of the Lamberticeras lamberti Zone (Z. Lewy, personal commun.) with a possible Upper Bathonian to Upper Callovian uncon- formity. ABBREVIATIONS AMNH American Museum of Natural History, Department of Invertebrates GSI Geological Survey of Israel, Paleontolo- gy Division USNM_ United States National Museum of Nat- AMERICAN MUSEUM NOVITATES NO. 2908 Fig. 2. Goliathyris lewyi, new species. A, B. Lateral, ventral views, holotype GSI M726la. x 1.1. ural History, Department of Paleobiol- ogy, Smithsonian Institution ACKNOWLEDGMENTS We thank Drs. C. H. C. Brunton, British Museum (Natural History), London, G. A. Cooper, National Museum of Natural His- tory, Washington, D.C., and Z. Lewy, Pa- leontology Division, Geological Survey of Is- rael, Jerusalem, for discussion, comments, and critical review of the manuscript. Feld- man is grateful to Dr. Y. Druckman, Head, Stratigraphy, Mapping and Oil Division, Geological Survey of Israel, for providing laboratory facilities and office space during his tenure as Visiting Scientist in Israel where a portion of the research for this paper was conducted. Feldman also acknowledges the assistance of Dr. F. Hirsch, Mapping Divi- sion, Geological Survey of Israel, for organ- izing field expeditions to the Sinai Peninsula and assisting in the interpretation of the com- plex regional stratigraphy. The research was funded by EARTH- WATCH and the Center for Field Research, Watertown, Mass., the Explorers Club Ex- ploration Fund, and the National Geographic Society under grant No. 2868-84 to Feldman. SYSTEMATIC PALEONTOLOGY SUBORDER TEREBRATULIDINA WAAGEN, 1883 SUPERFAMILY TEREBRATULACEA GRAY 1840 FAMILY UNCERTAIN Goliathyris, new genus TYPE SpeciEs: Goliathyris lewyi, new species. INCLUDED SPECIES: Type species only. GENERIC DIAGNOSIS: Extremely large, strongly sulcate, nonstrophic, permesothy- ridid with incurved dorsal umbo and massive zeilleriid beak. Broadly pentagonal in dorsal view. Hinge plates horizontal, becoming con- cave toward floor of brachial valve. Cardinal process massive and bilobate. CHARACTER ANALYSIS: Based on a clado- gram (fig. 6) it is postulated that Goliathyris is more closely related to Trigonithyris than to either Dyscolia or Goniobrochus, all three genera belonging to the family Dyscoliidae (Cooper, 1983; Muir-Wood, 1965). In sup- port of this hypothesis two synapomorphies are presented (table 3): (1) noncapillate radial ornamentation, and (2) a massive cardinal process. Goniobrochus is more closely related to Dyscolia than either is to Goliathyris or 1988 FELDMAN AND OWEN: GOLIATHYRIS LEWYI 5 Fig. 3. Goliathyris lewyi, new species. A, B, C. Posterior, anterior, dorsal views, holotype GSI M726 la. x1.1. Trigonithyris based on the following two syn- apomorphies: (1) ventrally directed hinge plates and (2) square loops. Dyscolia differs from Goliathyris in its sulcate anterior com- missure, permesothyridid pedicle foramen, and broadly pentagonal outline, all of which are considered plesiomorphic characters at this (generic) level of analysis. Cooper (1983) noted that Trigonithyris does not belong with the Dyscoliidae because of its well-developed outer hinge plates. Goliathyris resembles a zeilleriid externally in its beak and pedicle foramen (symplesiomorphies) but differs from the family Zeileriidae in having no dental plates or median septum. Based on analysis of the following characters the authors be- lieve that the present classification (Cooper, 1983; Muir-Wood, 1965) in which Dyscolia, Goniobrochus, and Trigonithyris are placed in the family Dyscoliidae must be reconsid- ered. Clearly, the four genera do not share any apomorphic characters. Additional data, such as the exact morphology of the loop in Trigonithyris and Goliathyris, will provide information in support or rejection of the proposed cladogram (fig. 6). Characters used in the above analysis and their phylogenetic significance are described below. 1. Dental plates: The character is absent in all four genera but present in the outgroup. This gives us no information regarding re- lationships among the genera. 2. Hinge plates: The character (ventrally 6 AMERICAN MUSEUM NOVITATES NO. 2908 Fig. 4. Goliathyris lewyi, new species. A, B. Lateral, dorsal views, paratype GSI M7261b. x1. directed hinge plates) is found in the out- group, Dyscolia, and Goniobrochus and thus represents a plesiomorphy at that level of analysis while its alternative character (con- cave hinge plates directed toward the brachial valve floor) represents a synapomorphy. 3. Cardinal process: The character is ple- siomorphic in Dyscolia (found also in the out- group) but synapomorphic in Trigonithyris and Goliathyris. 4. Ornamentation: The character (non- capillate radial ornamentation) is found in the outgroup, Trigonithyris, and Goliathyris, and thus represents a plesiomorphy at that level of analysis while its alternative char- acter (zigzag capillae) represents a synapo- morphy. 5. The loop: The character is found in Dys- colia and Goniobrochus and not in the out- group, thus representing a synapomorphy uniting the two genera; it is unknown in 77i- gonithyris and Goliathyris. If a zeilleriid loop is eventually found in these genera it would represent a plesiomorphy. Fig. 5. One of the auxiliary criteria of phylogenetic apomorphy recognized by Hennig (1966) for distinguishing apomorphic from plesio- morphic characters is used in the character analysis discussed above. The criterion used here is that of geological character precedence which, according to Wiley (1981), rests on the rule that when one character is found en- tirely in the geologically older members of a monophyletic group while the alternative ho- mology is found only in the geologically younger members of the same monophyletic group, then the older homolog is the plesio- morphic character. In accordance with the view that stratigraphic position not be con- sidered as a priori evidence for character evaluation (e.g., Schaeffer et al., 1972), cor- relation between stratigraphic position and relative primitiveness is now being consid- ered. The sister group Trigonithyris-Goli- athyris is represented only by Jurassic forms (Muir-Wood, 1965; this paper) and its genera share noncapillate radial ornamentation (character 4, above) postulated as plesio- ~ Transverse serial sections of Goliathyris lewyi, new species. GSI M7261b. Due to the nature of the matrix it was impossible to accurately record distances of serial sections from beak. However, the distance from the last transverse serial section to the umbo is 38 mm. Abbreviations: BHT = broken hinge tooth; CP = cardinal process; DP = descending processes; FHTr = flattened hinge trough; HP = 1988 FELDMAN AND OWEN: GOLIATHYRIS LEWYI 7 5 cm hinge plates; HT = hinge tooth; HTr = hinge trough; M = myophragm (= euseptoidum). Note how hinge plates change from almost horizontal to slightly concavoconvex, thicken, and point toward the floor of the brachial valve. 8 AMERICAN MUSEUM NOVITATES ZEILLERIIDAE TRIGONITHYRIS GOLIATHYRIS DYSCOLIA GONIOBROCHUS Fig. 6. A phylogenetic hypothesis of relation- ships among Jurassic, Pliocene, and Recent tera- bratulid brachiopods. See text for discussion. Filled boxes = apomorphic condition; empty boxes = plesiomorphic condition. morphic since it is also found in the outgroup (Zeilleriidae). Zigzag capillae, found in the sister group Dyscolia-Goniobrochus, which ranges in age from Pliocene to Recent (Coo- per, 1983), is a postulated synapomorphy, thus supporting the hypothesis that the al- ternative homology is the apomorphic char- acter and is found entirely within the geolog- ically younger members of a monophyletic group. However, upon examination of char- acters 2 and 3, it is evident that when apply- NO. 2908 ing the criterion of geological character pre- cedence there is an incongruity in that the apomorphic homologies are found entirely within the geologically older group. Addi- tional data will aid in resolving this problem. ETYMOLOGY: Named after Goliath, the Philistine giant from Gath, slain by [King] David. Goliathyris lewyi, new species Figures 2—5 DIAGNOSIS: Only known species of the ge- nus; same as for genus. DESCRIPTION: The shell is extremely large (table 2, figs. 2—4), smooth, rostrate, non- strophic, and broadly pentagonal when viewed dorsally. A small, circular pedicle fo- ramen is present just posterior to a poorly defined beak ridge on the ventral umbo, re- sulting in a permesothyridid condition. The interarea is obscured by the incurved ventral umbo. A wide, moderately shallow, dorsal sulcus originating slightly anterior to the con- cealed dorsal umbo, widens and deepens an- teriorly. Although much of the ventral ex- terior is poorly preserved, it is evident that a strong ventral fold opposed the dorsal sul- cus. The height of the fold appears to have been significantly greater than the depth of the sulcus. The shell is concavoconvex with the ventral valve almost carinate when viewed posteriorly. The anterior commissure is not preserved but, based on shell morphology, a reconstruction seems to indicate that it is strongly sulcate. One lateral commissure is preserved, extending about two-thirds of the shell length and, although partially concealed by debris anteriorly, appears to be uniformly straight. The lateral slopes of the ventral valve are quite steep, while on the dorsal valve they flare out from the sulcus, rise dorsally, and descend at approximately a 90° angle to the lateral commissure. Although radial ornament is absent on the smooth shell, there are some distinct, irreg- ularly spaced growth lines on the anterior half of the specimens. INTERNAL CHARACTERS: The following de- scription has been obtained with great diffi- culty from a series of transverse serial sec- tions (fig. 5) made through the umbonal region of a silicified specimen from the type locality at Gebel El-Minshera, northern Sinai. TABLE 2 Measurements of Goliathyris lewyi Compared with Homeomorphic Species of Aulacothyris from the “European” and “Ethiopian” Faunal Provinces (all measurements in millimeters) Faunal Species (L) (W) (T) Locality province Age Goliathyris lewyi, sp. nov. GSI M726la 74.0* 61.2 49.5 Gebel EP JUR El-Minshera GSI M7261b 72.6* _ 43.5 Gebel EP JUR El-Minshera Aulacothyris jubaensis 21.9 19.8 10.2 Somaliland EP JUR (Weir, 1929, pl. IV, fig. 15. XI) A. somaliensis (Stefanini, 25.0 20.0 15.0 Somaliland EP JUR 1931, pl. VI, fig. 6. XI) A, resupinata USNM 88703 22.8 19.7 15.0 Morocco EP JUR A, impressa AMNH 1514a 19.4 16.5 10.5 Bavaria EUP JUR AMNH 1514b 15.2 13.5 8.3 Bavaria AMNH 1514c 16.0 14.8* 8.0 Bavaria EUP JUR A, carinata AMNH 1509a 21.7 19.4 12.8 Bavaria EUP JUR AMNH 1509b 19.7 17.1 11.1 Bavaria EUP JUR AMNH 1509c 21.8 19.0 13.2 Bavaria EUP JUR A, pala AMNH 1523a 16.6 12.1 10.0 Vils in the Tyrol EUP JUR AMNH 1523b 19.7 12.9 11.5 Vils in the Tyrol EUP JUR AMNH 1523c 16.9 12.0 9.5 Vils in the Tyrol EUP JUR A. bernardina USNM 30949a 18.6 16.5 10.7 Villers-sur-mer, EUP JUR France USNM 30949b 15.8 13.9 8.3 Villers-sur-mer, EUP JUR France Aulacothyris sp. USNM 89083 21.5 16.3 11.2 England EUP JUR A. blakei USNM 104730a 14.6 13.2 7.9 England EUP JUR USNM 104730b 12.8 12.9 Tt England EUP JUR USNM 104730c 12.0 11.9 6.6 England EUP JUR A, meriani USNM 66932a 27.8 19.1 13.8 England EUP JUR USNM 66932b 21.7 16.3 12.7 England EUP JUR A. gregalis AMNH 19303a 10.2 10.8 6.1 Sarajevo, Bosnia EUP TRI AMNH 19303b 9.0 7.5 4.9 Sarajevo, Bosnia EUP TRI AMNH 19303c 9.4 9.3 4.8 Sarajevo, Bosnia EUP TRI A. angusta AMNH 1932a 7.0 6.4 4.0 Germany EUP TRI AMNH 1932b 71 7.1 4.4 Germany EUP TRI * Damaged. 10 AMERICAN MUSEUM NOVITATES NO. 2908 TABLE 3 Data Used to Analyze the Phylogenetic Relationships of Four Genera of Terebratulid Brachiopods Apomorphies are in italics. Outgroup Character (Zeilleriidae) Trigonithyris 1. Dental plates Present Absent 2. Hinge plates Ventrally direct- Concave toward ed brachial valve floor 3. Cardinal process Small Massive 4. Ornamentation Noncapillate Noncapillate 5. Loop Long Goliathyris Dyscolia Goniobrochus Absent Absent Absent Concave toward Ventrally direct- Ventrally direct- brachial valve ed ed floor Massive Small Absent Noncapillate Zigzag capillae Zigzag capillae Unknown Square Square . Unknown The umbonal cavity of the ventral valve is elongate-oval in transverse outline in the ear- ly stages and is partially filled with dense cal- lous. The cavity gradually develops a more triangular outline, meeting and articulating early with the dorsal valve. A massive, bi- lobate cardinal process develops and persists until the strong, peglike hinge teeth are well inserted into the dorsal sockets. Horizontal, elongate hinge plates, showing little or no dif- ferentiation from the inner socket ridges, de- velop a ventrally convex transverse outline and are deflected dorsally. No evidence was obtained of crural bases and all that remains of the obviously broken brachial loop is seen as two subparallel, in- wardly curving descending processes (or pos- sibly crural processes). Due to difficulty in sectioning and incompleteness of the shell, the last transverse serial section (fig. 5) was taken at 38 mm from the beak. ETYMOLOGY: The species is named for Dr. Zeev Lewy, Paleontology Division, Geolog- ical Survey of Israel, Jerusalem, in recogni- tion of his valuable contributions to the Me- sozoic paleontology of the Middle East. Types: The holoptype (GSI M7621a) and paratype (GSI M7621b) have been placed in repository in the paleontological collection of the Geological Survey of Israel, Jerusalem. OCCURRENCE AND AGE: Upper Callovian, Lamberticeras lamberti Zone, Gebel El- Minshera, northern Sinai (latitude and lon- gitude: 30°19’N, 33°43’E; Israel grid coordi- nates 9690/0219). REMARKS: This genus internally resembles Trigonithyris described by Muir-Wood (1935: 131) from the ?Argovian of British Somali- land. At the time, she suggested that the genus Trigonithyris was allied to Pygope and Pygites but later (Muir-Wood, 1965: H807) assigned it to the family Dyscolliidae, giving its strati- graphic position as Upper Jurassic (?Oxford- ian). The transverse serial sections of the type species Trigonithyris eruduwensis given by Muir-Wood (1935: 132) were produced be- fore the invention of the Croft serial grinding apparatus. Although poor by modern stan- dards, her sections clearly indicate the well- developed cardinal process and the almost horizontal, ventrally convex, and extensive hinge plates which show little or no differ- entiation from the inner socket ridges— mor- phological characters which distinguish this genus from other terebratulid genera so far examined from the ‘“‘Ethiopian’’ Faunal Province. Goliathyris lewyi has slightly con- cave to horizontal hinge plates (see fig. 5) which are very similar to Trigonithyris eru- duwensis Muir-Wood. However, G. lewyi is sulcate while 7. eruduwensis is rectimargin- ate with an erect beak and large pedicle fo- ramen. In Goliathyris lewyi, new species, the inner socket ridges appear to be more extensive and the cardinal process considerably more de- veloped than is seen in the serial sections of Muir-Wood’s genus. This may be due to the difference in relative size of the two speci- mens representing the species G. /ewyi and T. eruduwensis. Muir-Wood’s serial sections were from a small, adult form, whereas those of the very large G. /Jewyi specimen are of an old individual showing some evidence of ge- rontic thickening of the valves and cardina- lia. Although in its external morphology, es- 1988 pecially its zeilleriid beak, Goliathyris lewyi is suggestive of some terebratulidae, such as Rugitela impressa (Von Buch) and R. ber- nardina (D’Orbigny), it differs from these species in its less lobate outline and deeper sulcus. R. impressa has marked mesothyridid beak ridges and a distinct median septum in the dorsal valve, both of which are absent in G. lewyi. R. bernardina (D’Orbigny) also dif- fers in this way from G. /ewyi and has shal- lower valves, an acuminate to subpynform outline, and a shallower sulcus. Weir (1929) described a new terebratulid species as Aulacothyris jubaensis Weir (which is also figured in Stefanini, 1931), from the Juba Limestone of Dakatch, Somaliland, ranging in age from Callovian to Corralian. Although assigned to the terebratellacean ge- nus Aulacothyris, Weir’s species jubaensis shows little of the generic characters associ- ated with that genus apart from a similarity in general morphological outline, as is seen in the original description: Shell obovate-subpentagonal in outline, somewhat acuminate towards the anterior ex- tremity, which is regularly rounded and not re- entrant. In the region of greatest width, which is situated rather nearer to the posterior than to the anterior extremity of the shell, the lateral margins are subangular. Dorsal valve sulcate, the depression becoming more pronounced to- wards the anterior end; lateral surfaces of the dorsal valve convex, depressed to the level of the commissures. Ventral valve deep, the keel somewhat acutely rounded. Beak short, thick, closely pressed to the dorsal umbo, concealing the symphytium. Weir (op. cit.) contended that this species was closely related to “‘A. curvifrons” a species which is currently assigned to the terebratulid genus Pseudoglossothyris. In general outline it has much in common with Goliathyris lew- yi, new species, but differs from this species in its more acuminate anterior, having the dorsal valve more depressed toward the lat- eral margins and the pedicle foramen closer to the dorsal umbo concealing the symphy- tium. Also, its point of greatest width 1s near the posterior extremity and the lateral margin subangular. DISCUSSION: Brachiopods from the “Ethi- opian” Faunal Province are in need of study from both a taxonomic and paleobiogeo- FELDMAN AND OWEN: GOLIATHYRIS LEWYI 11 graphic point of view. Muir-Wood’s (1934, 1935) studies on Middle Jurassic brachio- pods, and especially those from the “‘Ethio- pian’? Faunal Province, need extensive re- vision before significant paleobiogeographic analysis can be undertaken. Evidence from East Africa suggests that there was a general faunal gradient in a north-south direction. In Callovian times the bivalve Neocrassina uni- lateralis was a southerly occurring species which was replaced northward by Neocras- sina scytalis. These changes are believed to be partly temperature controlled and are sup- ported by the occurrence of coral-Diceras de- posits in the Middle Jurassic of Ethiopia but not further south. The distribution of brachiopods follows these trends. An increased diversity of species occurs in the Callovian of Somaliland com- pared with beds of the same age from Tan- zania while the brachiopods of Kutch, India, are somewhat intermediate in morphology. Further study of the Jurassic brachiopods of northern Sinai, East Africa, Madagascar, and India will aid in reconstructing the early his- tory of the “Ethiopian” Faunal Province since the breakup of Gondwanaland. Since north- ern Sinai is situated at the tip of this province, the Sinai faunas are likely to include species belonging to the area within the equatorial Tethyan Realm which serves as a link be- tween the European faunas and those of Afro- Indian origin. REFERENCES CITED Ager, Derek V. 1973. Mesozoic Brachiopoda. In A. Hallam (ed.), Atlas of paleobiogeography, pp. 413-436. Amsterdam: Elsevier. Arkell, W. J. 1952. Jurassic ammonites from Jebel Tu- waygq, central Arabia. Philos. Trans. R. Soc. London, ser. B, 236: 241-313. 1956. Jurassic geology of the world. Edin- burgh and London: Oliver and Boyd, 806 pp. Cooper, G. Arthur 1983. The Terebratulacea (Brachiopoda), Triassic to Recent: a study of the bra- chidia (loops). Smithson. Contrib. Pa- leobiol., 50: 1-445. Hennig, Willi 1966. Phylogenetic systematics. Urbana: Univ. of Illinois Press, 263 pp. 12 AMERICAN MUSEUM NOVITATES Imlay, Ralph 1970. Some Jurassic ammonites from central Saudi Arabia. U.S. Geol. Surv. Profess. Pap. 643-D: D1-D17. King, William 1850. A monograph of the Permian fossils of England. Palaeontogr. Soc., Mon., 3: 1- 258. Kitchin, F. L. 1912. Palaeontological work; England and Wales: Geol. Survey of Great Britain and Museum Pract. Geology, Mem., Summ. of Prog. for 1911: 59-60. Muir-Wood, Helen M. 1934. 1935. 1965. On the internal structure of some Me- sozoic Brachiopoda. Philos. Trans. R. Soc. London, ser. B, 223: 511-567. The Mesozoic palaeontology of British Somaliland. Pt. 7, Jurassic Brachiopo- da, pp. 75-147, pls. 8-13. London: Government of the Somaliland Protec- torate. Mesozoic and Cenozoic Terebratulidi- na. In R. C. Moore (ed.), Treatise on invertebrate paleontology part H, Brachiopoda, pp. H762—-H816. Law- NO. 2908 rence, Kans.: Geological Society of America and Univ. Kansas Press. Schaeffer, B., M. K. Hecht, and N. Eldredge 1972. Paleontology and phylogeny. Evol. Biol., 6: 31-46. Stefanini, Guiseppe 1931. Werr, J. 1925. 1929. Echinodermi, Vermi, Briozoi e Bra- chiopodi del Giuralias della Somalia. Palaeontographia Italica, 32: 81-141. Brachiopoda, Lamellibranchiata, Gas- tropoda and Belemnites. Jn The collec- tion of fossils and rocks from Somali- land made by Mssrs. B. K. N. Wyllie and W. R. Smellie, Monogr. Geol. Dept. Hunterian Mus., Glasgow Univ., 1(6): 79-110, pls. 11-14. Jurassic fossils from Jubaland, East Af- rica, collected by V. J. Glenday, and the Jurassic geology of Somaliland III. Monogr. Geol. Dept. Hunterian Mus., Glasgow Univ., 3: 63 pp., pls. 1-5. Wiley, E. O. 1981. Phylogenetics: the theory and practice of phylogenetic systematics. New York: Wiley, 439 pp. Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates, Bulletin, and Anthropological Papers published during the last five years are available free of charge. Address orders to: American Museum of Natural History Library, Department D, Central Park West at 79th St., New York, N.Y. 10024.