Talexirhynchia, a New Rhynchonellid Genus from the Jurassic Ethiopian Province of Jordan

Talexirhynchia, a new rhynchonellid genus from the Jurassic Ethiopian Province of Jordan

Howard R. Feldman, Mena Schemm- Gregory, Mark A. Wilson & Fayez Ahmad

Paläontologische Zeitschrift Scientific Contributions to Palaeontology

ISSN 0031-0220

Paläontol Z DOI 10.1007/s12542-013-0216-y

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Pala¨ontol Z DOI 10.1007/s12542-013-0216-y

RESEARCH PAPER

Talexirhynchia, a new rhynchonellid genus from the Jurassic Ethiopian Province of Jordan

Howard R. Feldman • Mena Schemm-Gregory • Mark A. Wilson • Fayez Ahmad

Received: 30 May 2013 / Accepted: 28 November 2013 Ó Springer-Verlag Berlin Heidelberg 2013

Abstract A new genus and species of a rhynchonellide scattered on a limy substrate, such as shells and rocks, brachiopod from the Jurassic of Jordan, Talexirhynchia could have served as an attachment site for juveniles. With kadishi gen. et sp. nov., is described. The specimens were increasing growth, the loss of the pedicle and a semi- collected from the Mughanniyya Formation (Callovian) of infaunal position resulted in an increasingly incurved Wadi Zarqa from alternating claystones, siltstones, and ventral umbo that concealed the foramen. marly limestones with minor dolomite, dolomitic limestone, and coquinas that represent the upper part of the Keywords Talexirhynchia Á Rhynchonellida Á Jurassic sequence in Jordan. The environment of deposition Brachiopoda Á Middle Jurassic Á Ethiopian Province Á was neritic; food supply and light were unlikely to have Jordan been limiting factors. The specimens are related to Ethio- pian-Somali taxa and are consistent with the endemism that Kurzfassung Eine neue rhynchonellide Brachiopoden- characterizes the rhynchonellide brachiopod faunas of the Gattung und––Art, Talexirhynchia kadishi gen. et sp. nov., Jurassic Ethiopian Province. Specimens of Talexirhynchia aus dem Jura von Jordan wird beschrieben. Die Exemplare lived with the umbo in an upright position directed toward wurden in der Mughanniyya Formation (Callovium) von the seafloor or with the dorsal valve slightly above the Wadi Zarqa gesammelt und kommen in abwechselnden ventral valve. Juveniles were attached to the seafloor by the Lagen von Tonsteinen, Siltsteinen und mergeligen Kalken pedicle; carbonate shell material as well as other debris mit geringen Anteilen an Dolomiten, dolomitischen Kalken und Coquinas vor, die die ju¨ngsten jurassischen Ablager- ungen Jordaniens repra¨sentieren. Die Schichten wurden in M. Schemm-Gregory: Deceased. neritischer Fazies abgelagert, die wahrschein reich an Howard R. Feldman: Biology Department, Lander College for Nahrungsangebot und Licht durchflutet war. Die gesam- Women, The Anna Ruth and Mark Hasten School, A Division of melten Exemplare sind mit a¨thiopischen-somalischen Taxa Touro College, New York, NY 10023, USA. verwandt und spiegeln den Endemismus wieder, der die

H. R. Feldman (&) M. A. Wilson Division of Paleontology (Invertebrates), American Museum of Department of Geology, The College of Wooster, 944 College Natural History, New York, NY 10024, USA Mall, Wooster, OH 44691, USA e-mail: [email protected] e-mail: [email protected]

M. Schemm-Gregory F. Ahmad Centro de Geocieˆncias e Departamento de Cieˆncias da Terra, Department of Earth and Environmental Sciences, Faculty of Universidade de Coimbra, Largo Marqueˆs de Pombal, Natural Resources and Environment, The Hashemite University, 3000-272 Coimbra, Portugal P.O. Box 150459, 13115 Zarqa, Jordan e-mail: [email protected] M. Schemm-Gregory Museu Geolo´gico, Laborato´rio Nacional de Energia e Geologia (LNEG), Rua Academia das Cieˆncias, 19, 1200-003 Lisbon, Portugal

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H. R. Feldman et al. rhynchonellide Brachiopodenfauna der jurassischen Africogryphaea costellata, Homomya hortulana, and Pho- A¨ thiopischen Provininz charakterisiert. Vertreter von ladomya lirata. Dating of the Jordanian formations is partly Talexirhynchia leben in senkrechter Position mit der Dor- based on ammonite zonation (for example, the Bull- salklappe gering ho¨her als die Ventralklappe und dem atimorphites bullatus Zone that occurs in the Mughanniyya Wirbel zum Meeresboden hin gerichtet. Juvenile Exemp- Formation) and partly upon molluscan assemblages (Af- lare waren mit dem Stiel an Schalenmaterial und anderem ricogryphaea costellata–Eligmus asiaticus Zone from the Schuttdebris, z.B. Schalen oder Steine, der auf dem Hamam Formation) where ammonites are sparse. Upon Meeresboden verteilt vorlag, festgehaftet. Mit zuneh- completion of the taxonomic studies of the Jordanian bra- mender Gro¨ße a¨nderte sich die Lebensweise in eine semi- chiopods, we will be able to correlate the Middle Jurassic infaunale Position, die zu einem Verlust des Stieles und strata across the Dead Sea Rift (DSR). For example, the eines sta¨ndig weiter gebogenen ventralen Wirbels bis hin Callovian Mughanniyya Formation in the Zarqa region of zu vollsta¨ndigen Bedeckung des Stiellochs, fu¨hrte. northwest Jordan, east of the DSR, will be compared with the Zohar and Matmor formations of southern Israel and Schlu¨sselwo¨rter Talexirhynchia Á Rhynchonellida Á Gebel Minshera, Egypt, west of the DSR, as well as with the Brachiopoda Á Mittel-Jura Á A¨ thiopische Provinz Á Arroussiah Formation of Gebel El-Maghara, northern Sinai, Jordanian Egypt. The Mughanniyya Formation is correlative with the lowermost part of the Zohar Formation in the Negev that is coeval with the lowermost part of the Arroussiah Formation Introduction in Sinai, Egypt.

The first geological studies in this region were completed by geologists exploring the region for petroleum (Picard Geographic and geologic setting and Hirsch 1987). However, fossils belonging to the Jurassic Ethiopian Province have been collected as early as The Levant, from where this material was collected, is the late nineteenth century from Saudi Arabia (Mollet part of the elevated platform terrain of the Arabian 1871), as well as geographically widespread areas such as Nubian Shield that is covered by intermittent Palaeozoic Somalia (Muir-Wood 1935; Weir 1925, 1929), Egypt to Cenozoic sedimentary successions consisting mainly of (Douville´ 1916) and the Cutch, India (Kitchin 1900). The clastic units with marine carbonates increasing upward Jurassic macrofossil fauna from Jordan, mainly brachio- (Rybakov and Segev 2005). At the onset of the Mesozoic pods and molluscs, collected in this period was later this area consisted of broad sea-marginal flats, analogous described by Cox (1925) and Muir-Wood (1925). Begin- to modern sabkhas, in which evaporites and dolomites are ning in the mid twentieth century brachiopod workers have found, whereas in the Early Jurassic, emergence led to studied these faunas from a more modern point of view subaerial exposure that was accompanied by extensive (Cooper 1989; Dubar 1967; Farag and Gatinaud 1960a, b; freshwater runoff and subaerial weathering (Goldberg and Feldman et al. 2001). Brachiopods and related faunas from Friedman 1974). Shallow and marginal shelf environ- northern Sinai have been studied by Feldman (1987), ments interrupted by lagoons resulting in the deposition of Feldman and Owen (1988), Feldman et al. (1982) and thick, partly calcareous sandstones that were overlain by Hirsch (1979) and those from Jordan by Feldman et al. thick, partly gypsiferous carbonates, marls, and sandy (2012) and are currently under study. In general, the bra- marls that can be observed as a result of subsequent chiopods from the Ethiopian Province typically show a subsidence and faulting of the Dead Sea Rift (Basha high degree of endemism typified by genera such as 1980). Daghanirhynchia, Somalirhynchia, Somalithyris, Biheni- The Jurassic strata samples crop out in northwestern thyris and Striithyris, but despite the work by Muir-Wood Jordan along the western part of Wadi Zarqa beginning (1925, 1935), Hegab (1988, 1989, 1991a, b, 1992, 1993), near the old Jerash Bridge and extending westward to Deir- and Cooper’s (1989) monograph, the Middle Jurassic bra- Alla, a distance of about 20 km (Fig. 1); toward the south chiopod fauna of this region is still poorly studied. the outcrop belt passes through Ain-Khuneizir, Subeihi, Bivalves from northwest Jordan have been collected and and Arda Road (Ahmad 2002a, b). The Jurassic succession will be used in paleoecologic studies to determine the nature decreases in thickness from the Zarqa River and Wadi Huni of the marine communities of the Middle Jurassic in the eastward toward the Zarqa Bridge and from there southeast region. Examples of abundant bivalves collected from the toward Suweileh-1 and Safra-1. In the Zarqa River vicinity study area in Jordan include: Gryphaeligmus jabbokensis, the estimated total thickness of Jurassic ranges between Africogryphaea costellata, Eligmus rollandi var. jabbok- 290 m (Wadi Al Azab) and 400 m (Old Jarash Bridge and ensis, Chlamys textoria, Gryphaeligmus jabbokensis, Deir Alla) (Khalil and Muneizel 1992; Bandel 1981). 123 Author's personal copy

Talexirhynchia, a new rhynchonellid genus

Fig. 1 Geological map of the study area showing the collecting locality (1)

According to the lithological subdivision of the study brachiopods from the same horizon as Talexirhynchia area (Khalil and Muneizel 1992), our material occurs in the indicated that the original shell material appears to be Callovian Mughanniyya Formation. The Mughanniyya unaltered. In the specimens studied, original shell and Formation (Fig. 2) is composed of alternating claystones, sediment matrix appear not to have been diagenetically siltstones, and marly limestones with minor dolomite, altered nor recrystallized. dolomitic limestone, and coquinas; it represents the upper Several hundred serial sections were prepared with a part of the Jurassic sequence in Jordan cropping out con- WOKO 50P grinding machine with slice-spacing of sistently below the overlaying Kurnub Sandstone Group. 100–200 lm. Acetate peels were used to record the mor- phologic information from each slice. These were subsequently digitized using a digital camera (Canon 300 D). Materials and methods Three-dimensional reconstruction methods are those of (Sutton et al. 2001, 2005), using the custom SPIERS All specimens are preserved as articulated shells. To study software suite for registration, virtual preparation, and the internal morphology, serial sections are required. interactive visualization. The digitized acetate peels were Computer tomographic scans of these specimens proved subsequently manually aligned with SPIERSalign and, in a useless as the composition of shell material and sediment second step, edited by deﬁning different masks for the matrix does not allow sufﬁcient contrast for adequate ventral shell dorsal shell and crura in SPIERSedit. The 3D images. Cathodoluminescence testing of rhynchonellide images are copied out of SPIERSview. Drawings of the

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Fig. 2 Lithostratigraphic column of the Tel el Dhahab section. Bed S-7 with Talexirhynchia kadishi gen. et sp. nov. indicated by the brachiopod symbol

peels were made with the help of a camera lucida. Mea- Order Rhynchonellida Kuhn 1949 surements were taken with a digital caliper and rounded to Superfamily Hemithiridoidea Rzhonsnitskaya 1956 0.1 mm or using freeware ImageJ 1.43 l. Photographed Family Tetrarhynchiidae Ager 1965 specimens were coated with ammonium chloride. Subfamily Tetrarhynchiinae Ager 1965 The material herein was collected during various ﬁeld trips under strict stratigraphic control that will allow future Type genus: Tetrarhynchia Buckman 1918. paleoecologic studies of the brachiopods and a comparison Diagnosis: Dorsibiconvex to convexoplane trilobate Tet- with other marine communities from the area (e.g., the rarhynchiidae; multicostate, sometimes with short smooth Negev and Sinai). The studied and illustrated material is stage posteriorly; uniplicate, with dorsal fold well deﬁned, housed in a repository in the American Museum of Natural moderately to strongly raised, often subcarinate; linguiform History, New York City (AMNH). The systematics follows extension distinct, trapeziform to subtriangular, beak small, the revised Treatise on Invertebrate Palaeontology (Part H, usually suberect to incurved; delthyrium typically small, no. 4) (Mancen˜ido et al. 2002). foramen usually not rimmed. Hinge plates subhorizontal to convex ventrally; dental plates variable, subparallel, con- Systematic palaeontology vergent or divergent ventrally, conspicuous septalium, Y-shaped to pit-like; lateral umbonal chambers subtrian- Phylum Brachiopoda Du´meril 1806 gular, and empty. Crura raduliform, usually in the form of Subphylum Rhynchonelliformea Williams et al. 1996 simple hooks, occasionally somewhat expanded distally Class Rhynchonellata Williams et al. 1996 (after Mancen˜ido et al. 2002: 1344–1345).

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Genera included: Tetrarhynchia Buckman 1918, Cym- very low dorsal median septum, narrow and shallow sep- atorhynchia Buckman 1918, Goniorhynchia Buckman talium and the orientation of crura distally oriented parallel 1918, Grandirhynchia Buckman 1918, Quadratirhynchia to the commissural plane instead of extending in an ante- Buckman 1918, Somalirhynchia Weir 1925, Dag- roventral direction perpendicular to the commissural plane, hanirhynchia Muir-Wood 1935,?Belbekella Moiseev justify the erection of a new genus (see description of 1939, Robustirhynchia Seifert 1963, Orlovirhynchia Dagys Talexirhynchia kadishi). Shi and Grant (1993) that 1968, Eoseptaliphoria Ching and Sun in Ching, Sun and describes and illustrates ‘‘raduliform’’ crura that are typical Rong 1976, Druganirhynchia Tchoumatschenko 1983, of the Tetrarhynchinae; however, the crura of T. kadishi do Baeorhynchia Cooper 1989, Deltarhynchia Cooper 1989, not resemble their illustrations. In fact, the crura of T. Echyrosia Cooper 1989, Pycnoria Cooper 1989, Kor- kadishi differ from many of the related Tetrarhynchonellid jakirhynchia Smirnova 1990, Pontaltorhynchia Owen and genera described and illustrated in Shi and Grant (1993) Rose 1997, Talexirhynchia n. gen. and Cooper (1989). Externally, Talexirhynchia most clo- sely resembles Daghanirhynchia Muir-Wood 1935 and Genus Talexirhynchia nov. Septirhynchia Muir-Wood 1935. The lack of the ventral Type species: Talexirhynchia kadishi gen. et sp. nov. by median septum, the dorsibiconvex shell, the shorter umbos, monotypy which are less incurved, and the simple costae argue for an assignment to Daghanirhynchia and not to Septirhynchia, Derivation of name: After Alexa and Talia Belowich for even though Talexirhynchia differs from all Tetrarhyn- their assistance and interest in paleontology. chiinae in its strongly convex dorsal valve. Talexirhynchia Diagnosis: Tetrarhynchonellids with strongly convex dor- differs from Daghanirhynchia in having two to four fewer sal valve, little secondary shell material in dorsal apical costae on the fold than does Daghanirhynchia, smaller region, faint dorsal myophragm, septalium narrow and dental plates, shorter crura, and a slightly longer dorsal shallow, sockets smooth not corrugated, strongly shortened median septum. The morphological differences between anterior margin and consistently three costae on fold. Crura the new genus, Daghanirhynchia and Septirhynchia are short, raduliform, and strongly curved distally in a ventral summarized in Table 1. position until becoming parallel to the commissural plane, Taxa of Tetrarhynchia Buckman 1918 are more laterally thus, separating T. kadishi from many other Tetrarhynch- expanded and show coarser ribs than Talexirhynchia. onellid genera such as Aalenirhynchia, Colpotoria, Dag- Baeorhynchia Cooper 1989 differs from Talexirhynchia in hanirhynchia, Eurysites, Goniorhynchia, Heteromychus, its smaller size, an additional two to three more costae on Isjuminella and Kallirhynchia. fold and flanks, a longer beak, a longer dorsal median septum, and well-developed deltidial plates. The latter are Species included: Presently only the type-species Tale- lacking in Talexirhynchia. Cymatorhynchia is almost xirhynchia kadishi gen. et sp. nov. equibiconvex and less globose than Talexirhynchia and has Discussion: The presence of accreted secondary shell more, finer, and angular costae. Specimens commonly have material in the ventral apical region (see secondary shell about six costae on the fold, five costae in the sulcus, and material surrounding lateral apical cavity in Fig. 4k), the about eight costae on the flanks. Its crura are longer and

Table 1 Morphological comparison of Talexirhynchia, Daghanirhynchia, and Septirhynchia Talexirhynchia n. gen. Daghanirhynchia Muir-Wood Septirhynchia Muir-Wood 1935 1935

Crura Short Long Long Dorsal median septum Long, low, thin Short, low, thin Long, high, thin Ventral median septum Absent Absent Present Transverse outline Subelliptic in ventral–dorsal Subcircular to transverse Subelliptic in ventral–dorsal direction subquadrate direction Secondary shell material in apical Ventral valve None to much in both valves Much in both valves region Stratigraphic distribution Bathonian–Callovian Callovian Callovian–Kimmeridgian Geographic distribution Jordan Morocco, Tunisia, Jordan, Tunisia, Somalia, Ethiopia, Ethiopia, Somalia, Kenya, Kenya, Egypt (Sinai), Syria, Egypt (Sinai), Arabia, Israel, Oman, ?Iran, ?Madagascar, Syria, China, India ?Lebanon

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Fig. 3 Talexirhynchia kadishi gen. et sp. nov. 1a–e, 2–3, original size and oblique posterolateral (1m) views of external dorsal apical region, (91); 1f–o, magnified (92). 1. AMNH FI-75482, Tel el-Dhahab ventral (1m) and lateral (1o) views a three dimensional reconstruction section, Wadi Zarqa, Northwest Jordan, Bed S-7, Mughanniyya of the crura in the sectioned specimen. 2. AMNH FI-75483, Tel el- Formation, Callovian (upper Middle Jurassic). Ventral (1a), dorsal Dhahab section, Wadi Zarqa, Northwest Jordan, Bed S-7, Mug- (1b), lateral (1c), posterior (1d), and anterior (1e) views of articulated hanniyya Formation, Callovian (upper Middle Jurassic). Ventral (2a), specimen, internal view of apical region with (1f) and without (1g) dorsal (2b), lateral (2c), posterior (2d), and anterior (2e) views of crura, anterior view of internal ventral shell with (1h) and without (1i) articulated shell. 3. AMNH FI-75484, holotype, Tel el-Dhahab crura, dorsal view of external ventral apical region without (1j) and section, Wadi Zarqa, Northwest Jordan, Bed S-7, Mughanniyya with (1k) crura (note that although the crura are not morphologically Formation, Callovian (upper Middle Jurassic). Ventral (3a), dorsal part of the ventral valve we placed them in this position in order to (3b), lateral (3c), posterior (3d), and anterior (3e) views of articulated illustrate where they would be in the living specimen), ventral (1l) shell more strongly curved than in Talexirhynchia. Deltarhyn- foramen, and short dental plates. Goniorhynchia Buckman chia Cooper 1989 differs from Talexirhynchia in having an 1918 is less globose than Talexirhynchia and has finer and equibiconvex shell, a higher and longer dorsal median angular costae, of which five are on the fold, four in the septum, thick deltidial plates, a hypothyrid foramen and sulcus, and about eight on the flanks. Its specimens are long dental plates, whereas Talexirhynchia is characterized more widely transverse than representatives of Tale- by a strong dorsibiconvex curvature, a shorter and lower xirhynchia, the foramen is hypothyrid to submesothyrid, dorsal median septum, no deltidial plates, a mesothyrid whereas the foramen in Talexirhynchia is mesothyrid.

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Talexirhynchia kadishi sp. nov. posterior end of the umbo slightly anterior to the hinge line. The fold broadly rounded in cross section and strongly See Figs. 3, 4. elevated above the dorsal valve on the anterior half of the Etymology: After Alan Kadish, President of Touro Col- shell. Sparse accretion of secondary shell material in the lege, New York, USA. umbonal chamber of the ventral valve; however, lateral Holotype: Holotype, articulated specimen housed in the apical cavities are almost completely filled with it American Museum of Natural History AMNH F1-75484. (Fig. 4g–n). Posterior part of the dental plates thickened Tel el-Dhahab section, Wadi Zarqa, Northwest Jordan; Bed due to the accretion of calcareous callosity, anterior parts S-7, Mughanniyya Formation, Callovian (upper Middle of the dental plates are fine. Dental plates short and ending Jurassic). in small and knob-like teeth that are curved in an antero- lateral direction (Figs. 3-1j, 4j). Ventral median septum Type locality: Wadi Zarqa, northwest Jordan. lacking. Secondary shell material developed in the lateral Type horizon: Bed S-7, Mughanniyya Formation apical cavities but lacking in the central apical region. (Callovian). The dorsal median septum is fine and low, extending as a low ridge anteriorly in the posterior third of the shell. Stratigraphic and geographic distribution: See type Septalium is very small, narrow and shallow, poorly locality and type horizon. developed, however, according to Shi and Grant (1993: Fig. 13), small and incomplete septalial plates are often Diagnosis: Medium sized Talexirhynchia with shortened, developed, which do not reach the bottom of the shell stubby anterior margin and three costae on fold, strongly (Figs. 3-1f, g, 4g–i). Dental sockets are small and rounded convex dorsal valve, faint dorsal myophragm, narrow and in cross section (Fig. 4j). Inner socket walls are not curved shallow septalium, smooth sockets, short, raduliform crura over the socket ridges. Crural bases are thin, extending strongly curved distally in a ventral position until becom- from the hinge plate parallel to the commissural plane and ing parallel to the commissural plane. giving rise to very short, raduliform crura (Figs. 3-1k, n–o, Material: Eight articulated shells AMNH FI-75482-75487. 4k–m). Posterior ends of the crura are oriented perpen- (see Table 2 for measurements). dicular to the commissural plane (Fig. 4n). In other similar Description: Medium sized shells with subtriangular out- genera the crura are oriented with their ends in an anterior- line and geniculate dorsibiconvex in longitudinal section. ventral position rather than strictly ventrally; in other Dorsal valve globular, ventral valve almost flat (Fig. 3-1a–c). words they are perpendicular to the commissural plane. Shells thick in posterior region and very thin at the Discussion: Specimens of Pycnoria are equibiconvex and anterior margin, especially on the dorsal valves; anterior more strongly transverse in outline than Talexirhynchia. part of the shell slightly abraded (Figs. 3, 4-m, o, p). Both The anterior commissure of the fold and sulcus is more valves covered by medium size costae, which are rounded pronounced than in the new genus. Pycnoria shows finer to angular in cross section and separated by angular to costae with 2–3 more on the fold and sulcus than Tale- rounded furrows of same size as costae in depth and width. xirhynchia. The dental plates in Pycnoria are long and In the anterior half of the specimens, where the shell thick, but short and thin in Talexirhynchia. Rubustirhyn- becomes thinner, costae are impressed to the internal sur- chia shells are almost transversely semielliptical in outline; face. Each ventral flank bears five to six costae, dorsal costae are lower and less pronounced. The ventral beak flanks show three to four costae that weaken and fade extends much further posteriorly than the dorsal beak and laterally. Costae in sulcus and on fold are of same size as is not incurved. The teeth are long in Rubustirhynchia and costae on flanks. In rare cases costae on fold are coarser short in Talexirhynchia. Daghanirhynchia macfadyeni and costae in sulcus gently thinner than costae on flanks. bears 4–5 costae on the fold and has corrugated dental Costae on fold and sulci are not intercalated nor do they sockets. bifurcate. The sulcus bears two, whereas the fold bears three costae. Ventral beak incurved, slightly larger than dorsal beak Paleo-biogeographic implications but barely overlapping dorsal valve. Pedicle foramen hid- den by the incurvature of the dorsal beak, mesothyrid in 3D The brachiopods described herein belong to the Jurassic reconstruction (Fig. 3-1j). Ventral and dorsal planareas are Ethiopian Province, which has a wide geographical extent not visible. Anterior margin of shells are strongly unipli- ranging from east to North Africa including Somalia, cate. The sulcus and fold conspicuous. Sulcus low and Kenya, Tanzania, Saudi Arabia, the Levant, Egypt, Tuni- almost flat in cross section (Fig. 3-1a–3e) originating at the sia, and the Maghreb (Hirsch et al. 1998). The taxonomy of

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Fig. 4 Serial sections of Talexirhynchia kadishi gen. et sp. nov. posterior end of the shell (as abraded shell, c crus, cb crural base, dms AMNH FI-75482, Tel el-Dhahab section, Wadi Zarqa, Northwest dorsal median septum, dp dental plate, ds dental socket, dv dorsal Jordan, Bed S-7, Mughanniyya Formation, Callovian (upper Middle valve, lac lateral apical cavity, ssm secondary shell material, t tooth, Jurassic). All ﬁgures magniﬁed 92.0. Sectioning perpendicular to vv ventral valve) commissural plane, sectioning distances were measured in mm from

Table 2 Measurements of the material studied the brachiopod faunas here is complex due to the large Specimen Length/ Width/ Thickness/ degree of homeomorphy. Cooper (1989) comments that mm mm mm most earlier studies of these faunas were made without the adoption of serial sectioning and even the brachiopods AMNH-FI 75482* 21.0 20.7 20.1 identiﬁed with Buckman’s names in Muir-Wood’s (1935) AMNH-FI 75483 24.2 est 23.9 26.0 est monograph in the absence of interior details must be sus- AMNH-FI 75484 holotype 25.7 22.4 24.8 pect and indeed all are queried by their author. The Ethi- AMNH-FI 75485 28.1 25.4 23.8 opian Province has been recognized from the Early Jurassic AMNH-FI 75486 22.7 23.1 16.4 until the mid and possibly the end of the Cretaceous by the AMNH-FI 75487 20.7 18.3 17.1 presence of endemic taxa at the species, genus, and family Sectioned specimen is marked by the asterisk level (Feldman et al. 2001). The present fauna from Wadi

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Zarqa lies at the northernmost part of the Ethiopian Prov- tissue probably prevented additional clogging if sediment ince and may, therefore, be related to species of the did manage to seep between the valves. The strongly southern Tethyan margin to the west. Upon completion of a geniculate dorsibiconvex curvature of the shells and the systematic revision of the brachiopods from northwest elevated fold helped separate the inhalant and exhalant Jordan presently under study, we will be able to more currents that are maximized if the anterior margin of the accurately define faunal- and province-realm boundaries. sulcus is the highest point of the shell (Rudwick 1970; The taxonomic information recorded as a result of this Alexander 1999a, b; Savazzi 1999). This is comparable to study will add to our understanding of the biogeographic the chimney effect described by Vogel (1975) for spiriferid history of the Ethiopian Province and help to interpret the brachiopods, namely, oriented almost vertically with the structure and paleoecology of its marine communities. umbo down and the anterior commissure directed away from the sediment–water interface. The accretion of secondary shell material in the apical ventral valve and the Paleoecology lack of it in the apical dorsal region in Talexirhynchia supports the idea that the shell was oriented and also sta- Specimens of Talexirhynchia are found with other bilized in a living position with the dorsal valve above the rhynchonellid taxa (Feldman et al. 2012) and other inver- ventral valve, in which the tongue of the sulcus is in the tebrates. The environment is interpreted as a carbonate highest position. The large number of bottom dwelling facies with well-oxygenated water and high nutrient con- individuals in this assemblage required a reproductive tent, which may explain the high faunal diversity (see Wolf survival strategy. The strong curvature of the dorsal valve 1973). Specimens of Talexirhynchia compare favorably in provides Talexirhynchia with a relatively large body terms of morphology and functional morphology with chamber. We do not believe Talexirhynchia used an shells of Septirhynchia. It can be inferred from this that internal filtering system as proposed for spiriferids in a they also probably lived with the umbo directed toward the nutrient-poor environment (Schemm-Gregory 2010); seafloor in an upright position or with the dorsal valve instead we propose it had a storage place for long tentacles above the ventral valve. Juveniles were attached to the to provide a larger filtering area for the brachiopod, giving seafloor by the pedicle that atrophied as growth progressed. it an advantage over other rhynchonellids of similar size. In They could have been attached to various substrates such Recent brachiopods the ratio between body chamber and as shell debris and rocks. With increasing growth and free area is no longer used. The largest Recent brachiopods, pedicle loss, we propose a semi-infaunal life position as fist-size terebratulids from New Zealand, have an internal result of an increasingly incurved ventral umbo that con- loop that supports the lophophore; however, terebratulids cealed the foramen. Rudwick (1961) discussed the pre- represent another order that can hardly be compared to the sence of a pedicle consisting of a ramifying byssus-like rhynchonellids. We are not aware of any such strong dor- structure for brachiopods living in fine grained sediments sibiconvex rhynchonellids that could be used for compar- according to an observation on Recent Chlidonophora Dall ison and argumentation about our interpretation of the 1903 and Terebratulina D’Orbigny 1847a, b. The byssus- ecology of Talexirhynchia. like structure is also interpreted for large winged Paleozoic brachiopods with incurved umbos and could be a synapo- Acknowledgments Feldman would like to acknowledge the Touro morphic feature in Mesozoic brachiopods. However, we College Faculty Research Fund for supporting this project and thank the members of its peer review panel. We thank Jann Thompson and argue for the loss of the pedicle. The thickness of the apical Daniel Levin (National Museum of Natural History/Smithsonian shell and the coarse costae helped to stabilize the bra- Institution, Washington DC, USA) and Lee Davies (Natural History chiopod’s position within the sediment (Feldman 1987; Museum, London, UK) for providing access to the brachiopod col- Schemm-Gregory 2010). The thin anterior prevented the lection. Many thanks are due to Juliane Eberhardt and Erika Scheller- Wagner (Senckenberg Forschungsinstitut and Naturmuseum, Frank- brachiopod from further sinking into the sediment. The furt am Main, Germany) for technical help. The visit of M. S.-G. to semi-infaunal living position has been described as a mode the American Museum of Natural History, New York, was supported of life by various authors for brachiopods since the Pal- by a Lerner Grey Award. M. S.-G. thanks Russell Garwood (Man- aeozoic (Rudwick 1961, 1970; Ziegler et al. 1996; Ager chester University, UK) and Mark Sutton (Imperial College, London, UK) for instructions, help, and endless patience in solving SPIERS 1967; Grant 1971: 317, fig. 1; Westbroek et al. 1975: 374, problems. text-fig. 4; Richardson 1981; Schemm-Gregory and Jansen 2007; Zambito and Schemm-Gregory 2013). The overlap of the lateral dorsal valve above the ventral valve (Fig. 4n–p) References permits gaping of the valves without allowing mud to seep through the buried posterior portion of the commis- Ager, D. V. A. 1965. Mesozoic and Cenozoic Rhynchonellacea. In sure (Feldman 1987; Mancen˜ido and Walley 1979). Soft Treatise on Invertebrate Paleontology, Pt. H Brachiopoda 2, ed. 123 Author's personal copy

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