01 - Gourvennec.indd 1. INTRODUCTION The Taurus Mountains are one of the major tectonic units of southern Turkey, and many Palaeozoic outcrops have been recorded in this area, where the succession ranges from Cambrian to Miocene in age with many unconformities. The Devonian series show a regressive tendency following the transgressive sequence of the Silurian (see GEDIK, 1988, for a general overview of the Devonian of Turkey). The studied section is located NE of Kozan, on the road between Kozan and Feke, along the edge of Kilgen Lake, in SE Turkey, eastern part of the Taurids (Fig. 1). From East to West, the suc- cession includes the Ordovician series (Seydisehir Fm., Kozan Fm., Sortepe Fm. and Halevikdere Fm.) followed by Silurian black shales and limestones. The upper part of the latter, consisting of limestones lying unconforma- bly on the rest of the formation and underlying the Mes- ozoic formations, has yielded a fauna mainly consisting of brachiopods and corals. Although this succession was formerly considered as entirely Silurian, the brachiopod fauna collected in this upper part indicates a Late Devo- nian age, probably Early Frasnian. The age of the series is thus reassessed on this basis. Only the brachiopods are described here; study of the corals is in progress and they will be described in a future publication. Upper Devonian Brachiopods from Eastern Taurus (Turkey) Rémy GOURVENNEC 2. SYSTEMATIC PALAEONTOLOGY Note on the terminology The abbreviations used in the text and tables are the fol- lowing: L – length measured from the hinge to the ante- rior commissure; Lt – total length including the beak; W – maximum width; Wh – width of the hinge line; Ws – width of the sulcus or fold at the anterior commissure; Hd and Hv – height (i.e. depth) of the dorsal and ventral valves respectively; H – total height (i.e. thickness) of the shell; Ncf and Ncs – number of ribs (or costae) on one flank and sulcus or fold respectively; Ncp – number of parietal ribs. All measurements are in mm. The material is housed in the Laboratoire de Paléon- tologie de Brest, France (numbers LPB). Order Strophomenida ÖPIK, 1934 Superfamily Strophomenoidea KING, 1846 Family Douvillinidae CASTER, 1939 Genus Douvillina OEHLERT, 1887 Type species: Orthis dutertrei MURCHISON, 1840, p. 253. Douvillina sp. Pl. 1, Fig. 1 Material One shell fragment showing the ornament; another fragment showing the ventral muscle field. LPB 12400. Kozan section, level PK66 C. Description The shell has a low, planar, vertically striated interarea and a denticulate hinge. The delthyrium is triangular, partly filled by the basal thickening of the myophragm. The ornament is parvicostellate with well marked radial costae separated by wide, flat interspaces bearing fine costellae (2–3 costellae between primary costae), and tending to become stronger anteriorwards. Very fine concentric growth lines are also present on weakly expressed “rugae”. In the ventral valve, fine radial striae resulting from the alignment of numerous closely spaced small pits occur. They are separated by radial grooves, some of them deeper, mimicking the external ornament. The Geologia Croatica 59/1 1–17 1 Fig. 4 Tabs. 2 Pls. ZAGREB 2006 Key words: Brachiopods, Upper Devonian, Gondwa- na, Cimmerian Belt, Turkey. UMR 6538 “Domaines Océaniques”, Université de Bretagne Occidentale. UFR Sciences & Techniques. 6, Avenue Le Gorgeu, CS 93837, F-29238 Brest Cedex 3, France; e-mail: remy.gourvennec@univ-brest.fr Abstract During a field expedition in Taurus (SE Turkey) in order to study the Lower Palaeozoic series, a small brachiopod fauna was collect- ed in the Kozan area in beds of supposed Silurian age. After closer examination, the age of this fauna is clearly Upper Devonian, prob- ably Lower Frasnian, and this implies a reassessment of the age of the series. This assemblage also shows close affinities with similar North Gondwanan faunas described in Iran and Afghanistan in beds of the same age. 2 Geologia Croatica 59/1 muscle field is elevated, giving rise to a transverse, con- cave platform clearly delimited by an elevated ridge. It is crossed by a strong but relatively low median myo- phragm that does not run beyond the anterior edge of the muscle field. Discussion The internal features and ornament are consistent with an assignment of this species to Douvillina OEHLERT, 1887. D. ferquensis (RIGAUX, 1872) shows a similar ornament and arrangement of the internal pits related to the external ornament, but it differs in a longer myo- phragm that crosses the anterior margin of the mus- cle field. The ornament of some other species (e.g. D. aronovae LJASCHENKO, 1952 in LJASCHENKO, 1959) evokes the ornament of the present species, but due to the lack of material, it is difficult to make solid comparisons. These two species are known in Frasnian succes- sions of Western Europe and the Russian Platform respectively. Order Productida SARYTCHEVA & SOKOLSKAJA, 1959 Suborder Productidina WAAGEN, 1883 Superfamily Productoidea GRAY, 1840 Family Productellidae, SCHUCHERT, 1929 Genus Productella HALL, 1867 Type species: Productus subaculeatus MURCHISON, 1840, p. 255. Productella sp. cf. subaculeata (MURCHISON, 1840) Pl. 1, Figs. 2–5 Material Four almost complete variably damaged specimens, and two partially exfoliated ventral valves; LPB 12401 – LPB 12402. Kozan section, levels PK66 A and C. Description Small to medium size (means: L=18.3 mm; W=21.6 mm; H=8.7 mm), strongly concavo–convex with vari- able outline (rounded subquadratic to slightly trans- verse), with a hinge line slightly shorter than maximum width. The areas are discrete, very low, but distinguishable on most specimens and longitudinally striate; the angle between them varies from 90 to 130°. The delthyrium is broad, low, with no trace of deltidial cover. All the umbos are damaged, except on one specimen that shows no trace of an attachment cicatrix. Chilidium lacking. The shell bears fine, concentric sinuous growth lines and coarser, sinuous rugae which are stronger on the postero–lateral parts of the shell. The latter are some- what more conspicuous on the dorsal valve surface. They are generally low except on one specimen where they are well developed on the entire anterior half of the shell on both valves, and particularly well marked on the dorsal one. The spines, present on the ventral valve only, are long, hollow, numerous, recumbent, and distributed along the rugae in alternate rows resulting in an irregu- Fig. 1 Location map and lithostratigraphic column of the Kozan section, SE Turkey. The fauna described in this paper was sampled from the top of the series (level PK 66). 3Gourvennec: Upper Devonian Brachiopods from Eastern Taurus (Turkey) lar staggered implantation. However, due to the irregu- larity of the rugae, they often appear as randomly dis- tributed, particularly on abraded shells. They can be locally more or less aligned radially, rarely with pos- terior slightly elongate ridges. No spine was observed on the dorsal valve where they are replaced by small dimples approximately distributed in the same way as spines on the opposite valve. Due to lack of internal molds, the internal structures were not observed in detail. Nevertheless, two shells, partially exfoliated in their apical part, show a bifid cardinal process with high, divergent lobes and small sockets with short, rounded, slightly internally recurved teeth. Discussion The lack of ribs and cicatrix attachment, absence of spines in the dorsal valve, and the presence of teeth plead in favour of an assignment to Productella HALL, 1867. The closely related genus Spinulicosta NAL- IVKIN, 1937 shows a tendency to anterior ribbing, which is not the case here, and its spines generally originate from elongate spine bases. Donalosia LAZA- REV, 1989 presents a similar external aspect, but in Donalosia the spines are stronger and more numerous in the postero–lateral region, unlike the present form. Praewaagenoconcha SOKOLSKAJA, 1948 has a row of stronger median spines on the ventral valve, and Eostrophalosia STAINBROOK, 1943 has spines on the dorsal valve. Among the species belonging to the genus, P. sub- aculeata (MURCHISON, 1840) seems to be the better receptacle, although the spines are generally described as randomly distributed in this species, which is not exactly the case here, and less numerous (after the original figure in MURCHISON, 1840, pl. 2, fig. 9). As established by BROUSMICHE (1973) the species is highly variable in size, outline and ornament, but the different morphotypes described by this author always show relatively coarser spines than our specimens. We thus maintain this material in open nomenclature although the remaining characters are in good accord- ance with an assignment to P. subaculeata. Among oth- er close species, P. belanskii STAINBROOK, 1943 has a row of median stronger spines and elongated spine bases. P. subaculeata is known in Middle and Upper Devo- nian series from Europe, Asia, America, and Africa. Order Rhynchonellida KUHN, 1949 Superfamily Rhynchotrematoidea SCHUCHERT, 1913 Family Trigonirhynchiidae SCHMIDT, 1965 Genus Cupularostrum SARTENAER, 1961 Type species: Cupularostrum recticostatum SARTE- NAER, 1961, p. 2. Cupularostrum? sp. Material One complete crushed shell; LPB 12430. Kozan sec- tion, level PK66 C. Description This relatively large, moderately biconvex shell (Lt=27.5 mm; W=29 mm; H~15 mm) has a subcircu- lar outline. Sulcus and fold are not expressed or weak near the anterior margin that is damaged; the tongue was probably low. The ribs are numerous (Nc~50 over the entire shell), simple, angular, relatively strong, the median ones slightly stronger than lateral ones. The api- ces are smooth. The interior is unknown. Discussion Owing to the lack of information on the interior and the damaged commissure, this specimen is here assigned with doubt to Cupularostrum SARTENAER, 1961. It also presents some analogies with Comiotoechia LJAS- CHENKO, 1973 in its general aspect, but is larger, with coarser ribs. Genus Rhipidiorhynchus SARTENAER, 1966a Type species: Terebratula livonica von BUCH, 1834, p. 57. Rhipidiorhynchus cf. elburzensis (GAETANI, 1965) Pl. 1, Figs. 10–13 Material Only one crushed but complete specimen; LPB 12403. Kozan section, level PK66 B. Description Shell small (L=14 mm; W=14 mm), subquadratic, strongly inequivalve (Hd=8.5 mm; Hv~2.5 mm), with a maximum width located about mid-length. The maxi- mum height is located at the front. Ventral valve very low and concave (but this may be due to deformation). The beak is well developed, erect. The sulcus is clearly delimited, deep, wide (Ws/W=0.5 at front) with steep flanks, and originates in the poste- rior part of the valve, not very far from the apex. The tongue is high, trapezoidal. Dorsal valve very high, convex, with an obvious- ly delimited, high fold, originating in early stages of growth. Ornament of simple, rounded to subangular ribs, beginning at the apex and bearing very fine concentric growth lines (8 per mm). There are 3 ribs in the sulcus, 2 parietal ribs that do not reach the commissure, and 8–10 lateral ribs on each flank. The ribs on the fold are somewhat stronger than the others. Internal characters are unknown. 4 Geologia Croatica 59/1 Discussion This species is assigned to the genus Rhipidiorhyn- chus SARTENAER, 1966a on the basis of its ornament beginning at the apex, its sinal formula, the develop- ment and shape of the fold/sulcus, the erect beak, etc. Due to the lack of material, a specific assignment is unsure. This form is close to R. elburzensis (GAE- TANI, 1965) that was identified in the Frasnian of Iran and Afghanistan and differs by its larger size and high- er number of lateral ribs (but this is not significant in regard of the number of specimens available here). R. farsani BRICE, 1976 is also smaller with a different rib formula. Another closely related species, R. ferquensis (GOSSELET, 1887) is also discarded for the same rea- sons. The latter two are known in Frasnian successions, in Afghanistan and Europe respectively. Genus Cyphoterorhynchus SARTENAER, 1965 Type species: Uncinulus (Uncinulina) koraghensis REED, 1922, p. 40. Cyphoterorhynchus koraghensis (REED, 1922) Pl. 1, Figs. 19–23; Table 1 1922 Uncinulus (Uncinula) koraghensis sp. nov. REED, p. 40, pl. 7, figs. 10–22; pl. 8, fig. 1 1971 Cyphoterorhynchus koraghensis (REED, 1922) – BRICE, p. 47, pl. 3, figs. 1a–d, 6 a–d; text-fig. 13A (with a comprehensive synonymy). Material Seven articulated shells; LPB 12437– LPB 12439. Kozan section, levels PK66 A, C and D. Description Shell relatively large (means: Lt=22.3 mm; W=22.2 mm; H=19 mm) (Table 1), strongly dorsi–biconvex with subcircular to slightly transverse outline and vertical or subvertical lateral margins. The maximum height is located at mid-length, occasionally between mid-length and the anterior third of the shell. The sulcus and fold are poorly delimited and weak in the posterior part of the shell. They are also weak in the anterior half. The bottom of the sulcus is slightly convex. The tongue is high, vertical with subtrapezoidal outline. The anterior and lateral commissures are serrate. The ribs originate at the apex, are relatively strong, rounded to angular, never flattening anteriorly. There are usually 5 to 7 ribs in the sulcus and 1 (rarely 2) parietal rib(s) that invariably disappear anteriorly. Each flank bears 10 to 15 ribs (mean Ncf=12), the posterola- teral ones being very faint. Table 1 Measurements (in mm) of Cyphoterorhynchus koraghensis (REED, 1922) and C. arpaensis (ABRAMIAN, 1957) from the Kozan section, near Kilgen Lake, SE Turkey. Sample Lt W Hd Hv Ncs Ncp Ncf C. koraghensis LPB 12437 — — — — 4 1 >12 LPB 12438a 25.5 25.5 15 8.5 6 1 10 LPB 12438b 21.5 21.5 12 6 5 1 10–12 LPB 12438c 21.5 24 12 5.5 5 1 10 LPB 12439a 21 20.5 11 7 6 1 (2?) > 13 LPB 12439b 22 21 13 5.5 7 2 ≈15 LPB 12439c 22.5 20.5 13 5.5 7 1–2 14 Means 22.3 22.2 12.7 6.3 ≈6 ≈1 ≈12 C. arpaensis LPB 12431a 22.5 23 10 5 6 4 13 LPB 12431b 21.5 20 13 5 7–8 2 >18 LPB 12432 22 22 15 5 7 2 >17 LPB 12433a 19.5 20 11 6 7 3 >18 LPB 12433b 17 17 8 4 8 3 >13 LPB 12434a 27 22 12 7 ≈9 3 >19 LPB 12434b 19.5 19 12 9 ≈8 3 >13 LPB 12435a 25 23.5 14 7 9 2 >16 LPB 12435b 18 18 10 5 7 2 15 LPB 12436a 22 21.5 9 7 9 3 >12 LPB 12436b 16.5 18 8 7 8 3 >13 Means 21 20.4 11.1 6.1 ≈8 2–3 ≈16+ 5Gourvennec: Upper Devonian Brachiopods from Eastern Taurus (Turkey) Dorsal interior with septalium; other internal struc- tures unknown. Discussion See the discussion of C. arpaensis below. The species is known in Frasnian successions of Pakistan (Chitral), Afghanistan, and Iran. Cyphoterorhynchus arpaensis (ABRAMIAN, 1957) Pl. 1, Figs. 24–28; Table 1 1957 Camarotoechia radiata NAL. (in litt.) var. arpaen- sis var. nov. ABRAMIAN, p. 55, pl. 5, figs. 6 a–г; pl. 6, figs. 4 a–г. 1971 Cyphoterorhynchus arpaensis (ABRAMIAN) – BRICE, p. 52, pl. 3, figs. 2 a–c, 4 a–d; text-fig. 14 A (with a comprehensive synonymy). Material 14 specimens; LPB 12431 – LPB 12436. Kozan section, levels PK66 A, B, C, D. Description This species is very similar to the previous one and dif- fers mainly by the shape and number of ribs that are finer, more numerous, less angular and occasionally become flattened anteriorly, with a median groove (vis- ible on two specimens). There are 7 to 9 ribs in the sul- cus and 2 or 3 parietal ribs that disappear anteriorly in about 50 percent of the specimens. In one case, there are 4 parietal ribs. The number of lateral ribs is highly variable (13 to more than 19 on each flank) (Table 1). The deltidial plates are low and disjunct. Ventral interior with short, thick dental plates that converge ventrally. The muscle field is excavated and crossed by a short median myophragm and a few radial grooves. A strong thickening of the shell may be present, reducing the free part of the dental plates. Dorsal interior with a strong V-shaped septalium with a dorsally convex cover plate. The long, relatively high median septum is thick posteriorly and becomes abruptly thinner anteriorly. Crural bases subcircular in cross-section. The imprint of the anterior and posterior adductor scars is well marked and moderately excavat- ed. On two specimens the vascula are well developed in both valves, particularly the vascula lateralia. The vascula genitalia are distributed anterolaterally to the muscle field and consist of small tubercles more or less radially oriented and sometimes fused together, in the posterior third of the valve. Discussion In this species, as for the species described above, the variability seems very pronounced. It concerns the general shape, more or less transverse, sometimes sub- cuboid, the number of ribs and their strength, etc. This variability both in C. koraghensis and C. arpaensis has already been observed in material described in Afghani- stan (BRICE, 1971), Iran (SARTENAER, 1966b) or Chitral (REED, 1922). Although the extremes are eas- ily separated, the presence of transitional forms does not make it easy to distinguish the two species by their external aspect; the main criterion consists of the differ- ent sinal formulae and number of lateral ribs. Some of the intermediate morphs (with 2 parietal and few lateral ribs) may belong to the subspecies C. koraghensis inter- positus SARTENAER, 1966b which is a transitional form between the nominal species and C. arpaensis. The two species are assigned here to Cyphotero- rhynchus SARTENAER, 1965, following the general opinion of the authors in recent literature. Nevertheless, they also show some similarities with the genus Cupu- larostrum SARTENAER, 1961 such as trapezoidal tongue, serrate commissure, distinct dental plates more or less ventrally convergent and relatively long dorsal septum. C. arpaensis has been described from the Frasnian of Armenia, Transcaucasus, Iran and Afghanistan. Superfamily Pugnacoidea RZHONSNITSKAJA, 1956 Family Ladogiidae LJASCHENKO, 1973 Genus Ladogilina LJASCHENKO, 1973 Type species: Ladogilina rossica LJASCHENKO, 1973, p. 38. Ladogilina sp. 1 Pl.1, Figs. 6–9 ; Table 2 Material Three almost complete specimens and one crushed specimen; LPB 12426 – LPB 12428. Kozan section, levels PK66 A, B, C. Description Shell strongly inequivalve, large (maximum: L=38 mm; W=40 mm) (Table 2), with a rounded subtriangular outline. The apical angle varies from 100 to 110°. The maximum width is located near the anterior third of the shell. Ventral valve low, more or less flat in transverse section, a very shallow sulcus appearing only at mid- length. At the anterior edge, the tongue is very high and vertically erect, its basis occupying almost the entire width of the shell. The extremity (top) of the tongue is rounded. The beak is short, well marked, with an appar- ently hypothyrid foramen. Dorsal valve very high and regularly increasing in height up to the frontal commissure. In transverse sec- tion, the valve is regularly rounded (semicircular shape) with a fold becoming expressed only in the anterior third of the shell. 6 Geologia Croatica 59/1 The entire shell is covered by numerous fine, round- ed, simple ribs beginning at the apex, separated by narrow interspaces and regularly widening anteriorly. There are 8 ribs per 5 mm at 1.5 cm from the beak and 4 to 6 ribs per 5 mm at the antero–lateral commissure. The dorsal median septum is strong, particularly in its posterior part. It is long, reaching the midlength of the shell. The ventral interior features are unknown. Discussion Despite its subtriangular outline, this species is assigned to Ladogilina rather than Ladogia after the shape of its tongue (not acute as in Ladogia), rounded ribs and the rounded outline in transverse section. This species is close to Ladogilina persanica BRI- CE, 1999, but differs by its larger size. BRICE (1999 – p. 73) pointed out the existence in Afghanistan (Robat- e-Paï) of a large specimen close to L. persanica that “possibly belong[ed] to another species”; this specimen could be conspecific with the present one. L. rossica LJASCHENKO, 1973 and L. simensis (MARKOWSKY, in ELLERN et al., 1955) are also much smaller than the present species and have somewhat finer ribs (9 to 11 per 0.5 mm at 1.5 cm from the beak). Ladogilina sp. 2 cf. L. rossica LJASCHENKO, 1973 Pl. 1, Figs. 29–33; Table 2 Material Two complete specimens and 2 incomplete, crushed ones; LPB 12429. Kozan section, level PK66 E. Description Shell inequivalve, relatively large, but smaller than Ladogia sp. 1 (maximum: L=28 mm; W=31 mm) (Table 2), with a rounded subquadrangular outline. The apical angle is about 110 to 120°. The maximum width is located in the anterior half of the shell. Ventral valve lower than the dorsal one, rounded in transverse section. Sulcus almost lacking or very shal- low, appearing at mid-length. The tongue is widely rounded, not very high. The beak is short with a subme- sothyrid foramen. Dorsal valve high and regularly rounded in trans- verse section. Fold indistinct. Ribs numerous and fine (11 to 12 per 5 mm at 1.5 cm from the beak and 6 to 8 per 5 mm at the com- missure). The dorsal median septum is thick, particularly in its posterior part and becomes relatively thin anteriorly. It is long, reaching the mid-length of the shell. The dental plates are short, thin and vertical. The ventral muscle field is poorly impressed and marked by a few, very low radial ridges. Discussion This species is close to the former one but differs in its more quadratic outline and its smaller size, although it remains larger than previously known species of the genus. The ventral valve is rounded in cross section, in contrast with Ladogilina sp. 1 (where it is flattened); it is also proportionally higher than in Ladogilina sp. 1. Furthermore, the ribs are finer and more numerous. This characteristic is reminiscent of both L. rossica and L. simensis in which the number of costae in 5 mm at 1.5 cm from the beak is around 9 to 11. Ladogilina sp. 2 is very close to L. rossica in its external aspect and internal features; it differs only by its larger size. The genus Ladogilina is known in Russia (Urals and Timan), Eastern Iran (Kerman Province) and western Afghanistan. All the occurrences are from the Lower Frasnian (Asymmetricus Zone). Order Atrypida RZHONSNITSKAJA, 1960 Suborder Atrypidina MOORE, 1952 Superfamily Atrypoidea GILL, 1871 Family Atrypidae GILL, 1871 Subgenus Spinatrypa (Spinatrypa) STAINBROOK, 1951 Type species: Atrypa hystrix var. occidentalis HALL, 1858, p. 515. Table 2 Measurements (in mm) of the two species of Ladogilina from the Kozan section, near Kilgen Lake, SE Turkey. Sample L W Hd Hv Nc per 5 mm at Nc per 5 mm at 1.5 cm from beak commissure Ladogilina sp. 1 LPB 12426 38 39 20 8 8 4 LPB 12428a ≈30 ≈40 15 7 8 5 LPB 12428b 32 34 20 9 8 5 Ladogilina sp. 2 LPB 12429a 28 31 12 11 11 6 LPB 12429b 28 35 14 10 12 7 7Gourvennec: Upper Devonian Brachiopods from Eastern Taurus (Turkey) Spinatrypa (Spinatrypa) sp. e.g. S. longispina (RIGAUX, 1872) Pl. 1, Figs. 14–18; Table 3 Material Eight complete specimens and some separate valves; LPB 12404 – LPB 12409. Kozan section, levels PK 66A, B, C, D. Description Shell equibiconvex in juvenile growth stages becom- ing strongly dorsi–biconvex in adult stages (means of dorsal and ventral H respectively 6.9 and 4 mm – Table 3), slightly wider than long (means: L=19.6 mm; W=21.4 mm) with a hinge line shorter than the maxi- mum width. Sulcus and fold absent, only indicated by a slight ondulation of the anterior commissure in adult specimens. Ventral valve regularly convex; sulcus lacking in juveniles, becoming slightly developed in the immedi- ate vicinity of the anterior commissure in adults. Tongue very low and wide (exceeding 1/2 of shell width). A sin- gle gerontic specimen shows a relatively well defined sulcus and tongue. Umbo feebly prominent with a wide submesothyrid foramen. Interarea not developed. Dorsal valve more convex than the ventral one, reg- ularly postero–anteriorly curved. In transverse section, the valve is strongly convex in its median part, becom- ing concave in its postero–lateral extremities. The fold is absent. The ornament consists of coarse ribs (about 7 ribs per 10 mm near the commissure) increasing in number by dichotomy and intercalation. The growth lamellae are long (1 to 2 mm), and their anterior edge is sharply erect (with an angle reaching or exceeding 45°). They are covered with fine concentric growth lines (about 15 per mm). Spines are not preserved in situ, but some remains were observed in the sediment covering the external surface of one shell. Internal features unknown. Discussion From the general shape, ornament (ribs and growth lines) and despite the lack of information about its internal morphology, this form is undoubtedly related to the genus Spinatrypa STAINBROOK, 1951. Given the relative convexity of the valves (shell rather dorsibicon- vex) this form belongs to the subgenus S. (Spinatrypa). Among the numerous species assigned to the genus, this one is very close to S. longispina (RIGAUX, 1872) and differs only by its smaller size. The S. aspera group and S. curvirostra COPPER, 1967 have stronger, wider and less numerous ribs (4 ribs per 10 mm at the com- missure). The adult/gerontic specimens of the present species show some similarities with Atrypa nefedovae LJASCHENKO, 1950 in their strongly dorsiconvex profile, but differ by their smaller size and less numer- ous, coarser ribs. The only ascertained occurrence of S. longispina is in the upper part of the Ferques Formation (Boulonnais, N. France), in the Triangularis Zone (Frasnian). After GODEFROID (1988), the other mentioned occurrences (Belgium and Germany) are dubious or erroneous. Order Spiriferida WAAGEN, 1883 Suborder Spiriferidina WAAGEN, 1883 Superfamily Cyrtospiriferoidea TERMIER & TERMIER, 1949 Family Cyrtospiriferidae TERMIER & TERMIER, 1949 Genus Cyrtospirifer NALIVKIN, 1924 (in FREDERIKS) Type species: Spirifer verneuili MURCHISON, 1840, p. 252. Cyrtospirifer sp. aff. C. kermanensis BRICE, 1999 Pl. 2, Figs. 19–23 Material Nine more or less complete specimens and 3 ventral valves; LPB 12412 – LPB 12417a. Kozan section, lev- els PK66 A to E. Table 3 Measurements (in mm) of Spinatrypa sp. cf. S. longispina (RIGAUX, 1872) from the Kozan section, near Kilgen Lake, SE Turkey. Sample L W Hd Hv Nc (distance from the beak in mm) (5) (10) (15) (20) (25) (40) LPB 12404a 21.5 25.5 7.5 6.2 19 25 — — 33 — LPB 12406a 22 22 7.9 3.5 14? 22? ≈30 ≈36 — — LPB 12406b 14 15.5 4.6 3 ≈18 ≈30 30 — — — LPB 12407a 18.5 21.5 3.5 ≈14 ≈20 27 — — — LPB 12407b ≈11.5 ≈13 4 2.5 ≈20 ≈22 — — — — LPB 12407c 29.5 31.5 8.9 ≈7 18 22 — ≈32 50? — LPB 12407d (gerontic) 35.5 36.3 20 ≈5 16 24 28 36? 40? >50 LPB 12408a 12 13 3.5 2 14 21 24 — — — LPB 12408b 12 14.5 2.5 2.5 24 ≈40 — — — — Means 19.6 21.4 6.9 4 ≈17 ≈25 ≈28 ≈35 ≈41 — 8 Geologia Croatica 59/1 Description Shell small (means: L=12 mm; Lt=16 mm; W=22.3 mm without mucros), transverse with subtrapezoidal outline and slightly sinusoidal lateral commissures. The ven- tral valve is higher than the dorsal one (means: Hd=5.2 mm; Hv=7.5 mm). The shell is mucronate and the maxi- mum width is thus located at the hinge line. Shells with preserved mucros would probably reach a total width of 30 mm. When the mucros are broken, the cardinal extremities are subangular. The ventral interarea is low, apsacline, flat on its greater height, becoming abruptly curved in its upper part. It is clearly delimited (sharp angles at the transi- tion to the flanks) with subparallel edges, longitudinally and vertically striated, with finer vertical striation. The delthyrium is triangular and bounded by high, rarely preserved deltidial plates. The beak is strong, overhang- ing on the area, obscuring the top of the delthyrium in posterior view. The sulcus is deep, wide (~1/3 W at the anterior margin), and easily delimited by ribs stronger than any adjacent ones. It forms a wide open V-shaped or widely rounded section. The fold is variably expressed, gener- ally low, occasionally not protruding over the flanks, rounded in section. It is always clearly delimited by grooves, obviously deeper than adjacent ones. The tongue is rounded, variably developed, generally low. The entire shell is ribbed. The ribs on the sulcus/fold are narrower than those on the flanks, particularly in the vicinity of the anterior commissure. The ribs are sim- ple on the flanks (about 20 on each flank), the postero– lateral ones, near mucros, being very faint and poorly expressed. Their number increases by dichotomy (or intercalation?) on the sulcus/fold. These dichotomies/ intercalations occur relatively late (on the anterior half of the shell). There are generally 11 ribs (extremes: 10 and 12) on the sulcus/fold. The micro-ornament consists of fine, closely spaced concentric growth lines (about 7 per mm) and very fine radial capillae bearing densely distributed spine bases. Interior: the apical part of the shell is variably thick- ened. The dental plates are long (1/3 L), divergent, almost straight and extrasinal. A well developed del- thyrial plate is present. Ctenophoridium present; crural plates apparently lacking or obsolescent. The remain- ing internal structures have not been observed, due to recrystallization of the shells. Discussion The finely costate fold and sulcus, and the wide intera- rea with acute angles are consistent with assignment to the Cyrtospiriferinae. Cyrtospirifer NALIVKIN in FREDERIKS (1924) seems to be the best classification for this species although most of the representatives of this genus show a high, triangular interarea, which is not the case here. The general aspect of our species also recalls Geminisulcispirifer SARTENAER, 1982, but the latter has a characteristic median groove on the fold, intrasinal dental plates and bifurcations on lateral ribs. Some specimens of Sinospirifer sinensis GRABAU, 1931, particularly those with low interarea, are similar to our species but their mucros are basally wider, their sulcus shallower, the beak less curved and the number of lateral ribs is greater. Finally the most similar species is Cyrtospirifer ker- manensis BRICE, 1999, which differs from our species in its shallower, not so sharply limited sulcus and a gen- erally subquadratic outline. C. archiaci (MURCHISON, 1840) has a similar number of ribs, grooves bounding the fold and fine mucros, but it is more globose and elongate, with a subquadratic outline. C. disjunctus (SOWERBY, 1840) often considered as a synonym of C. verneuili (MURCHISON, 1840), shows fine mucros similar to those of our species, a low ventral interarea with subparallel edges and a fold clearly delimited by deep grooves, but it is much larger with a subquad- ratic outline. C. echinosus LJASCHENKO, 1958, is a transverse form with deep fold-bounding grooves, but its mucros are poorly developed, its ribs more numer- ous (on both flanks and sulcus) and its ventral interarea high and rather triangular. Another species, C. poljani- cus LJASCHENKO, 1954 in LJASCHENKO (1959) has fine mucros but a shallower, poorly delimited sul- cus and corresponding fold. C. kermanensis has been described from the Fras- nian of Iran. Cyrtospirifer sp. cf. C. verneuili (MURCHISON, 1840) Pl. 2, Figs. 5–7 Material Two complete, crushed specimens and 1 incomplete ventral valve; LPB 12417c and LPB 12425. Kozan sec- tion, levels PK66 A and E. Description These poorly preserved specimens are ventribiconvex, relatively large (maximum: L=27 mm, W=40 mm, H=24 mm) with a subquadrangular outline, wider than Eodmitria sp. (see description below). The ventral interarea is low (possibly owing to distortion) with a poorly prominent, more or less incurved beak. Cardi- nal extremities are acute with incipient mucronation. The sulcus and fold are clearly delimited but very shal- low/low, almost inconsistent, even in the proximity of the anterior commissure where the tongue is rather low. The sulcus has a flat V-shaped section; the fold has a rounded one. The ornament consists of very fine ribs over the entire shell. There are about 20 ribs on the fold/sulcus, where they increase in number by intercala- tion, and about 35 ribs on each flank on the larger spec- imen. They are finer than in C. sp. aff. C. kermanensis (see above) and similar in size to those of Eodmitria sp. (see below), but not flattened on top. The interior is unknown. 9Gourvennec: Upper Devonian Brachiopods from Eastern Taurus (Turkey) Discussion These specimens are assigned to Cyrtospirifer but owing to their poor preservation, a specific assignment remains uncertain. They differ from C. sp. aff. C. ker- manensis by their larger size, poorly developed fold and sulcus, fine ribbing and ventral interarea lacking sub- parallel edges. They differ from Eodmitria sp. by their transverse outline, more or less mucronate extremities and relatively coarser, not flattened ribs, but they show some affinities with some unpublished specimens of Eodmitria sp. from the lower Frasnian of Ferques (Bou- lonnais, France) presently under investigation (BRICE, pers. comm., 2004). Genus Eodmitria BRICE, 1982 Type species: Eodmitria supradisjuncta boloniensis BRICE, 1982, p. 578. Eodmitria sp. Pl. 2, Figs. 8–18; Table 4 Material 12 almost complete specimens and 1 isolated ventral valve; LPB 12417b – LPB 12424. Kozan section, levels PK66 A and C. Description Shell medium to large (means: L=24.8 mm; W=30.4 mm), slightly inequivalve (means: Hd=8.8 mm; Hv=11.9 mm) (Table 4), with variable outline: subquadratic weakly elongated, rarely slightly transverse (in adult stage). The maximum width is located around the midlength. The lateral commissures are widely round- ed, sometimes becoming slightly concave near the car- dinal extremities. The hinge line is very short (mean: Wh=19.1 mm), clearly shorter than the maximum width. Cardinal extremities subangular (widely obtuse angle) with traces of weak mucronation in some specimens. The ventral interarea is triangular, occasionally abruptly interrupted distally. It is well delimited, short, relatively high, curved and apsacline, with strong verti- cal striae. The delthyrium is triangular, lined by appar- ently low deltidial plates and deltidial slots. The beak is prominent, recurved on the interarea. Sulcus and fold very weak, becoming somewhat better expressed near the anterior commissure where a low, rounded tongue is developed. The sulcus is poor- ly delimited, the fold better delimited by two lateral grooves deeper and larger than adjacent ones. At the anterior margin, their width reaches or exceeds half of the maximum width (mean Ws=14.3 mm). In transverse section, the sulcus is widely rounded or slightly suban- gular, particularly in the juvenile stages. The entire shell is ribbed. The ribs are rounded with a flattened, or even depressed top, and separated by narrow, rounded to subangular interspaces. The ribs are generally simple on the flanks, with a few excep- tions, the distal ones being very faint. Their number lies between 30 and 38 per flank. Dichotomies, more rarely trichotomies, occur on the sulcus and fold, where the ribs are finer than the lateral ones. There are 13 to 27 ribs at the anterior edge of the sulcus/fold (mean=20.9). The microornament consists of very close prominent growth lines (4 to 5 per mm), with irregularly distribut- ed spine bases or pustules. Ventral interior: the dental plates are thin, straight, relatively long (about 1/4 L), slightly internally convex in section. The delthyrial plate is thin, and relatively long. There is no apical thickening of the shell. Muscle prints were not observed. Table 4 Measurements (in mm) of Eodmitria sp. from the Kozan section, near Kilgen Lake, SE Turkey. Sample L Lt W Wh Ws Hd Hv Ncf Ncs LPB 12418a 20 25 25 15 ≈10 6 10 ≈30 ≈13 LPB 12418b 23 28 27 18.5 14 9 15 30 20 LPB 12418c 20.5 27 26 12.5 ≈12 7 10 33 18 LPB 12419a 24.5 30.5 31 22 14 10 12 34 ≈20 LPB 12419b 26.5 32.5 32.5 27 ≈14 ≈7? ≈12 — — LPB 12421a 21 28.5 ≈31 21 ≈14 — ≈10 32 ≈21 LPB 12424a — — 31.5 16 — 10 11 35 20–22 LPB 12424b 34 39.5 38.5 23.5 19 11 13 37–38 ≈27 LPB 12424c 28 34 33 ≈18 15 10 13 ≈30 23 LPB 12424d 27 32 ≈34 21 15 9 13 37 20–21 LPB 12424e 28 34 34 19 16.5 9 14 >30 21–22 LPB 12424f 20.5 25.5 24.5 17 ≈12 8 10 33 19 LPB 12424g 24.5 31.5 ≈27 ≈18 ≈16 ≈9 ≈12 31 ≈26 Means 24.8 30.7 30.4 19.1 14.3 8.8 11.9 ≈33 ≈21 10 Geologia Croatica 59/1 Dorsal interior: the cardinal process is striated. The crural bases are long. Discussion The general outline of the shell, the weak, ill-defined sulcus and fold, the number of ribs and their shape, the presence of a well developed delthyrial plate, and the presence of crural plates, etc. indicate that this spe- cies belongs to Eodmitria BRICE, 1982. In the closely related genus Dichospirifer BRICE, 1971, the ribs are dichotomous on the entire shell and the delthyrial plate is rudimentary. The closely related genus Uchtospirifer LJASCHENKO, 1957 possesses a well developed del- thyrial cover and the sulcus and fold are clearly marked. In the original diagnosis, BRICE (1982 – p. 575) indi- cates that the shell is always mucronate in Eodmitria. Nevertheless, some species assigned to the genus since its erection show only a very slight mucronation. Despite its ressemblance to several species of the genus, the nomenclature of this species remains open because no known species possess this combination of characters. Eodmitria supradisjuncta (OBRUTCH- EV, 1917) differs from Eodmitria sp. in its thick den- tal and delthyrial plates and a thickened apical region. The subspecies E. supradisjuncta boloniensis BRICE, 1982 (type-species of the genus) also shows an apical thickening and has less numerous lateral ribs, a more transverse outline and a more pronounced tendency to mucronation. E.? seminoi (VERNEUIL, 1850) is also apically thickened, has intra- to subsinal dental plates and lacks crural bases. E. oblivialis SARTENAER, 1982 is larger, with a generally longer hinge line, and has stronger, less numerous ribs. It is possible that this species could be the same as that cited by TCHIHATCHEFF (1854) in the region of Feké (Anti-Taurus) under the name Spirifer seminoi VERNEUIL. The genus Eodmitria is known from the Lower Fra- snian of Western Europe (Asymmetricus Zone) and the Middle Frasnian from the Russian Platform (Timanicus and Triangularis Zones). Suborder Delthyridina IVANOVA, 1972 Superfamily Delthyridoidea PHILLIPS, 1841 Family Mucrospiriferidae BOUCOT, 1959 Genus Eleutherokomma CRICKMAY, 1950 Type species: Eleutherokomma hamiltoni CRICKMAY, 1950, p. 220. Eleutherokomma hamiltoni? CRICKMAY, 1950 Pl. 2, Figs. 1–4 Material Three ventral valves and one interior of a ventral valve; LPB 12410 – LPB 12411. Kozan section, levels PK66 B and C. Description Transverse shell with subtrapezoidal outline, later- ally developing very long mucros. The largest shell is 60 mm wide including mucros (that is about 30 mm without mucros) and 12 mm long. The area is relatively low, strongly apsacline, separated from the flanks by angular edges. The beak is barely prominent. The trian- gular delthyrium is bordered by low deltidial plates that probably join (?) under the umbo. The sulcus is wide (Ws=7.5 mm on the above men- tioned specimen), shallow, somewhat flared with a rounded section, and bears a faint median ridge. The ornament consists of 7 lateral rounded ribs per flank, separated by narrow subangular interspaces. The external pair is very faint but the internal pair (bounding sulcus) is clearly stronger than the others. Microorna- ment consists of wavy growth lamellae, the density of which increases anteriorly, and that bear very fine con- centric growth lines (8 per mm). Ventral interior with strong apical thickening embed- ding the apical structures: delthyrial ridge (rather than true delthyrial plate) and obsolescent dental plates. Teeth short, small, elliptical in outline. No trace of mus- cle scars observed. Dorsal valve unknown. Discussion After the general aspect of the shell, and particularly the long mucronations, this species can undoubtedly be assigned to Eleutherokomma CRICKMAY, 1950. The closest species is E. hamiltoni CRICKMAY, 1950, which presents a comparable number of ribs, with sul- cus bounding costae stronger than the lateral ones, and very long mucros. It differs in the presence of short dental plates that are partly free and thus visible, due to a less significant shell apical thickening. This variation could result from different environmental conditions. The other species of Eleutherokomma with few lateral ribs as in this species, have either short mucros or they are absent. E. hamiltoni is known from the Lower Frasnian of Canada (lower Asymmetricus Zone). 3. CONCLUSIONS Despite the fact that the nomenclature of many species described here remains open, due to the few specimens at hand, an estimation of the age can be done with rea- sonable certainty. At the generic level, this assemblage indicates a Late Devonian age: 4 genera have a Middle– Upper Devonian distribution, 2 are Upper Devonian and 3 are Frasnian (Douvillina, Eodmitria and Ladog- ilina), one of them (Ladogilina) being restricted to the Lower Frasnian. Otherwise, the presence of Spinatrypa precludes a Famennian age. This leaves no doubt about a Frasnian age and an Early Frasnian age is probable, 11Gourvennec: Upper Devonian Brachiopods from Eastern Taurus (Turkey) although Cyphoterorhynchus is mentioned from the Middle–Upper Frasnian. This is reinforced by an exam- ination at the species level: even if the specific assign- ments are not definitely stated, all the specific proximi- ties recognized here are with Frasnian species (e.g. R. elburzensis, C. koraghensis, S. longispina, E. hamiltoni, C. kermanensis, etc.). The association shows close affinities with those described from Iran and Afghanistan (BRICE, 1971, 1999) and also from the Russian Platform, Timan and Volgo–Uralian areas (LJASCHENKO, 1959, 1973). Several genera are common to these regions, and it is probable that the differences and/or uncertainties at the specific level could be clarified after complementary sampling in Turkey. It is clear that our material is insuf- ficient for a detailed study, but the obvious similarities in the fossil assemblages are troubling if not significant, particularly if we consider some genera with relatively restricted geographic distributions (e.g. Eleutherokom- ma and Ladogilina). Resemblances also occur at the morphological level, since several taxa from these dif- ferent regions share peculiar shell “habitus”: highly developed, globose ventral valves (Ladogia, Spina- trypa), sub-cuboid shells (Ripidiorhynchus, Cyphotero- rhynchus variants reminiscent of Hypothyridina) or the contrasted very short/very wide hinged spiriferids (Eod- mitria vs. Eleutherokomma). This morphologic analogy is probably related to similar environmental conditions. The faunal similarities between Iran, Afghani- stan, Pakistan and Armenia during the Upper Devo- nian have been already underlined by several authors (e.g. BROCK & YAZDI, 2000). The Devonian Turk- ish fauna indicates that the Taurids (at least the eastern part of the Taurids) should be included in this group of regions constituting the so-called “Cimmerian belt”, a suite of terranes located along the North Gondwanan margin. Furthermore, the presence of Eleutherokomma, also known in Iran, Mauritania (RACHEBOEUF et al., 2004), northern France and eastern Canada, reinforces the faunal evidence for a narrow oceanic separation between North Gondwana and Euramerica at this time, contrary to some reconstructions based mainly on pal- aeomagnetic data. Acknowledgements Many thanks to O. Monod (University of Orléans, France) who sampled and placed this material at my disposal. I am also grateful to the reviewers, U. Jansen and J. Sremac, for their helpful comments on the manu- script and J. Robson for the language review. 4. REFERENCES ABRAMIAN, M.S. 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(1840): On the physical structure and older stratified deposits of Devonshire.– In: SEDGWICK, 13Gourvennec: Upper Devonian Brachiopods from Eastern Taurus (Turkey) A. & MURCHISON, R.I. (eds.): On the distribution and classification of the older or Paleozoic deposits of the North of Germany and Belgium and their comparison with formations of the same age in British isles. Transac- tions of the Geological Society of London, 2nd series, 5/3, 633–703. STAINBROOK, M.A. (1943): Strophomenacea of the Cedar Valley limestone of Iowa.– Journal of Paleontology, 17, 39–59. STAINBROOK, M.A. (1951): Substitution for the pre-occup- ied brachiopod name Hystricina.– Journal of the Washing- ton Academy of Sciences, 41, 196. TCHIHATCHEFF, P. (1854): Dépôts paléozoïques de la Cap- padoce et du Bosphore [Paleozoic deposits of the Cappa- docia and Bosphorus – in French].– Bulletin de la Société Géologique de France, 2ième Série, 11, 402–416. VERNEUIL, E. de (1850): Notice sur la collection de roches recueillie en Asie par feu Hommaire de Hell, et sur les divers travaux exécutés pendant le cours de son voya- ge (In: VIQUESNEL, M.) [Notice on the rock collection sampled in Asia by the late Hommaire de Hell, and on various studies realized during the course of his trip – in French].– Bulletin de la Société Géologique de France, 2ième Série, 7, 491–514. Manuscript received April 6, 2005. Revised manuscript accepted November 11, 2005. 14 Geologia Croatica 59/1 PLATE 1 Fig. 1 Douvillina sp. Level PK66C. Interior of ventral valve, x2, LPB 12400. Figs. 2–5 Productella sp. cf. P. subaculeata (MURCHISON, 1840). Level PK66C. Dorsal, ventral, posterior and lateral views, x1.5, LPB 12402. Figs. 6–9 Ladogilina sp. 1. Level PK66A. Ventral, dorsal, posterior and lateral views, x1, LPB 12426. Figs. 10–13 Ripidiorhynchus sp. cf. R. elburzensis (GAETANI, 1965). Level PK66B. Lateral, dorsal, anterior and ventral views, x1.5, LPB 12403. Figs. 14–18 Spinatrypa (Spinatrypa) sp. e.g. S. longispina (RIGAUX, 1872). Level PK66C. Dorsal, ventral, ante- rior, posterior and lateral views, x1, LPB 12407. Figs. 19–23 Cyphoterorhynchus koraghensis (REED, 1922). Level PK66C. Dorsal, ventral, anterior, posterior and lateral views, x1, LPB 12438. Figs. 24–28 Cyphoterorhynchus arpaensis (ABRAMIAN, 1957). Level PK66C. Dorsal, ventral, anterior, posterior and lateral views, x1, LPB 12435. Figs. 29–33 Ladogilina sp. 2 cf. L. rossica LJASCHENKO, 1973. Level PK66E. 29–32: Dorsal, ventral, anterior and posterior views, x1, LPB 12429a; 33: posterior view of internal mould, x1.5, LPB 12429d. All specimens are from the Kozan section (Kilgen Lake, Eastern Taurus, Central Turkey). 15Gourvennec PLATE 1 1 2 3 4 5 6 7 8 9 10 11 12 14 15 13 16 18 17 23 19 20 21 24 25 26 29 30 31 27 33 32 28 22 16 Geologia Croatica 59/1 PLATE 2 Figs. 1–4 Eleutherokomma hamiltoni? CRICKMAY, 1950. Level PK66C. 1–3: ventral, anterior and lateral views, LPB 12411a; 4: interior of ventral valve, LPB 12411c. Figs. 5–7 Cyrtospirifer sp. cf. C.verneuili (MURCHISON, 1840). Level PK66E. 5–6: dorsal and ventral views; 7: microornament (x10), LPB 12417c. Figs. 8–12 Eodmitria sp. Level PK66A. Dorsal, ventral, anterior, posterior and lateral views, LPB 12418b. Fig. 13 Eodmitria sp. Level PK66A. Microornament (x10). LPB 12422. Figs. 14–18 Eodmitria sp. Level PK66C. Anterior, dorsal, ventral, posterior and lateral views, LPB 12424. Figs. 19–23 Cyrtospirifer sp. aff. C. kermanensis BRICE, 1999. Level PK66C. Dorsal, lateral, ventral, anterior and posterior views, LPB 12415. All specimens are from the Kozan section (Kilgen Lake, Eastern Taurus, Central Turkey) and x1.5, unless otherwise indicated. 17Gourvennec PLATE 2 1 2 3 4 8 7 65 9 10 11 12 13 14 15 16 17 19 20 21 18 22 23 18 Geologia Croatica 59/1