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Cretaceous Research 34 (2012) 257e267

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Cretaceous Research

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First report of Hadrosia Cooper, 1983 in South America and its biostratigraphical and palaeobiogeographical implications

Mena Schemm-Gregory a,b,*, Alexis Rojas-Briceño a, Pedro Patarroyo c, Carlos Jaramillo a a Smithsonian Tropical Research Institute, P.O. Box 0843-03092, Panama City, Panama b Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Paläozoologie II, D-60325 Frankfurt am Main, Germany c Departamento de Geociencias, Universidad Nacional de Colombia, Cr. 30 N. 45-03 or A. A. 14490, Bogotá, Colombia article info abstract

Article history: A new terebratulid species, Hadrosia gracilis, from the Lower Cretaceous Rosablanca Formation in Central Received 9 December 2010 Colombia is described. The new method of three-dimensional reconstructions of the internal morphology Accepted in revised form 9 November 2011 of this taxon results in the subjective synonymization of the Nerthebrochinae in favour of the Available online 17 November 2011 Sellithyridinae. The geographical distribution of species of Hadrosia in South America and Western Europe is a further argument for a direct pathway between these two regions during Early Cretaceous time. Keywords: Ó 2011 Elsevier Ltd. All rights reserved. Terebratulida Brachiopoda Rosablanca Formation Lower Cretaceous Colombia

1. Introduction description of the majority of the pre-Barremian fossil content using modern palaeontological techniques is still lacking. In the eastern Andes of Colombia there are extensive outcrops of This paper is one of a series on the Cretaceous brachiopods from a shallow marine, Lower Cretaceous sedimentary sequence consist- Colombia that we have carried out. Our data will help to establish ing of more than 1000 m of carbonate and siliciclastic rocks. Up to the history of Cretaceous brachiopod species, their evolution within now, most of these outcrops have only been described in the the Andean region, and their relationship to southern Tethyan framework of petroleum exploration (e.g., Cooper et al., 1995). faunas. The taxonomic information recorded as a result of these However, detailed studies of the lithology and certain fossil groups studies will elucidate the biogeographic history of the Andean with main focus on ammonites have been carried out during the past region and help to interpret the structure and palaeoecology of its half century (e.g., Bürgl, 1956; Etayo-Serna, 1964, 1968a, b, 1979; marine communities. Patarroyo, 2000a, b, 2004; Bogdanova and Hoedemaeker, 2004; Sharikadze et al., 2004). Further fossil groups considered are bivalves (e.g., Dietrich,1938; Villamil,1996), crustaceans (e.g., Feldmann et al., 2. Geological settings and previous research 1999; Feldmann and Villamil, 2002; Vega et al., 2007, 2010), vertebrates (Langston et al.,1955; Hernandéz-Camacho and De Porta, 2.1. Geological settings 1963; Páramo, 1994, 1997, 2000; Cadena and Gaffney, 2005; Ezcurra, 2009), and plants (Huertas, 1967, 1970, 1976; Waveren et al., 2002; The basal calcareous rocks of the Cretaceous in Colombia are Correa et al., 2010). With exception of testudines and decapod crus- present in the Middle Magdalena Valley, including the Villa de taceans (Cadena and Gaffney, 2005), however, a substantial Leyva area. They are represented mainly by the Rosablanca Formation (Morales et al., 1958; Etayo-Serna, 1968a, b; Rámon et al., 2001). The studied locality of the Rosablanca Formation is situated near the “la Fábrica” cave south of the town of Santa Soﬁa, northern * Corresponding author. Present address: Centro de Geosciências e Departamento Boyacá Province, central Colombia (Fig. 1). de Ciências da Terra da Universidade de Coimbra, Largo Marquês de Pombal The Rosablanca Formation is described as a black micritic and 3000-272, Portugal. Tel.: þ351 239 860 571; fax: þ351 239 860 501. almost pure limestone sequence with a small percentage of clastic E-mail addresses: [email protected], [email protected] (M. Schemm-Gregory), [email protected] (A. Rojas-Briceño), [email protected]. components leading into grainstones and evaporites (Morales et al., co (P. Patarroyo), [email protected] (C. Jaramillo). 1958). In the study area the Rosablanca Formation is conformably

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Fig. 1. Map of the study area with collecting locality indicated.

overlain by quartz siltstones of the Ritoque Formation (Etayo-Serna, 1968a, b; Ballesteros and Nivia, 1985; Patarroyo, 1997b, 2009). Cardozo and Ramirez (1985) described in detail the lithostratigraphy of the Rosablanca Formation in the Ayal Creek at Santa Sofía. At this section the formation consists of a lower sequence of 20 m of mudstones, muddy sandstones, and grainstones followed by 103 m of micritic, biomicritic and calcareous mudstones, calcareous sand- Fig. 2. Stratigraphic column for the Lower Cretaceous in Colombia. stones, and dolomitized micrites. Specimens of Hadrosia occur in beds of calcareous mudstones at the top of the Rosablanca Forma- central basin to the north where the Rosablanca Formation deposits tion; according to Cardozo and Ramirez (1985), between 10 and are younger (e.g., Morales et al., 1958; Patarroyo, 1997b). 12 m below the onset of the overlying Ritoque Formation (Fig. 2). The sedimentary environment of the Rosablanca Formation is 2.2. Studies on Colombian brachiopods interpreted as a broad carbonate platform (Morales et al., 1958; Rámon et al., 2001). The dark to medium grey colour of the beds The oldest brachiopods of Colombia were reported by bearing Hadrosia indicates lowered oxygen levels with a relatively Harrington and Kay (1951) from the CambrianeOrdovician deposits high supply of organic material (Pedersen and Calvert, 1990). in the Macarena Range without accurate stratigraphic location. Cardozo and Ramirez (1985) reported Chondrites and Thalassinoides Mojica et al. (1988) reported graptolites and some brachiopods in the Upper Rosablanca Formation in Santa Sofía, which argues for from the Middle Ordovician Hígado Formation in Tarqui, Huila local niches of more oxygenated conditions. They interpreted these Province. Bogotá (1982) and Thery et al. (1984) described strata as subtidal deposits adjacent to a tidal ﬂat environment. In his brachiopods and indeterminate impressions that we, in accordance study on Cretaceous bivalves from Zapatoca, Santander Province, with the authors, consider to be brachiopod remains in Ordovician central Colombia, Dietrich (1938) described the brachiopod genus sediments of the Colombian Amazon Basin. Sellithyris as frequently found in “calizas margosas” or muddy The most important work on Palaeozoic brachiopods is limestones, but gave no further sedimentological information. undoubtedly the monograph by Caster (1939) on brachiopods from According to new ammonite data near Villa de Leyva (e.g., the Devonian EmsianeEifelian Floresta Formation exposed near the Karakaschiceras Thieuloy, 1971, Raimondiceras Spath, 1924) strata of village of Floresta, central Colombia (see also Barret, 1988). the Rosablanca Formation are assigned to the Valanginian Stage. Recently, Janvier and Villarroel (2000) and Moreno (2004) However elsewhere, the uppermost part of the Rosablanca described lingulid brachiopods from this unit. Formation is referred to the Hauterivian (Haas, 1960; Guzman, Carboniferous brachiopods from Colombia, which are found in 1985) or HauterivianeBarremian (Patarroyo, 1997a) near Zapatoca strata of the Garzon, Quetame, and Santander Massif (Trumpy, 1943; or questionably lower Hauterivian (e.g., Acanthodiscus Uhlig, 1905) Olsson, 1956), have been studied by various authors: e.g., Gerth near Villa de Leyva, Boyacá Province. (1932), Kehrer (1933), Royo (1945), Stibane and Forero (1969),and The upper diachronous boundary of this lithostratigraphic unit Angiolini et al. (2003). Mojica et al. (1988) recorded brachiopods from is known because of a progressive transgressive event from the the same time interval, but without any speciﬁcdeterminations. Author's personal copy

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Mesozoic brachiopods from Colombia have been poorly studied Therefore, we suggest that Hadrosia gracilis sp. nov. is a possible index hitherto. Several authors (e.g., Geyer, 1973; Stibane, 1976; Sandy, fossil for the upper part of the Valanginian Stage in this region. 1990, 1991a, b) showed that some marine, almost unfossiliferous sediments of Palaeozoic to Jurassic age were misplaced for many 3.2. Palaeobiogeographical value years in the “red beds of Colombia”. Cretaceous brachiopods described so far include the terebratulids Arenaciarcula beaumonti Owing to the lack of modern palaeontological studies, (d’Archiac, 1847) and Gemmarcula cf. menardi (Lamarck, 1819), a consensus biogeographical interpretation of the Early Cretaceous stored in the d’Orbigny Collection in Paris, as well as Musculina aff. of Colombia is still lacking (Hoedemaeker, 2004). sanctaecrucis (Catzigras, 1948)(Guzman, 1985). Representatives of On a global scale, taxa of the Sellithyrididae have been used to Sellithyris sella (Sowerby, 1823) from Zapatoca were reported by interpret the palaeobiogeography and faunal relationships among d’Orbigny (1851), Karsten (1858, 1886), and Dietrich (1938) under Tethyan faunas during the Early Cretaceous (e.g., Sandy et al., 1995; the names Terebratula sella Sowerby, 1823 and Terebratula haueri Grãdinaru et al., 2006). One example is the cosmopolitan distribu- Klippstein, 1844. The most recent studies on brachiopods have been tion of the low latitude Tethyan genus Sellithyris which indicates the carried out by Rojas-Briceño and Patarroyo (2009, 2010); however, presence of faunal pathways in coincidence with the opening of the the data published so far are mostly preliminary. North Atlantic Ocean (Sandy, 1990, 1991a, b, 1997; Gaspard, 2005). Cenozoic brachiopods from Colombia are basically unknown. Manceñido (2002) also discussed faunal exchange and the rela- Preliminary reports consist of Recent terebratulids from the tionship between brachiopod faunas in northern South America and Caribbean Coast (Rojas-Briceño et al., 2009). Monospecific clusters Western Europe as a result of the opening of the South Atlantic of the discinid brachiopod Discradisca Stenzel, 1964 found in the Ocean; however, this fauna pathway occurred somewhat later. The Malaga Bay on the Pacific coast are currently under investigation faunal relationship between South America and North America and (Rojas-Briceño, unpublished data). with Europe, furthermore, is recognizable in coral and molluscan faunas (Von Der Osten, 1957). The occurrence of Hadrosia in France 3. Stratigraphy and palaeobiogeography of Hadrosia and Colombia confirms the hypothesis of a faunal migration way through the low latitude Tethys (Fig. 3), which was first discussed by 3.1. Biostratigraphic value Haas (1960). He described the similarity of Lower Cretaceous ammonites found in a locality about 100 km southeast of the locality Mesozoic brachiopods are characterized by strong homoplasy “Santa Sofía” in our study area to other Tethyan forms. On other and a decline in abundance and diversity towards the Cenozoic. hand, Etayo-Serna et al. (1976) argued that at the specific level the Their high intraspecific and intrageneric variability does not ease Hauterivian ammonites from Colombia are closely related to those their use as biostratigraphical and palaeobiogeographic tools. of Peru and at the generic level to Mexican forms. However, previous studies have shown that Cretaceous brachiopods are important for biostratigraphy and palaeobiogeography during this time interval (e.g., Grãdinaru et al., 2006; Lobatscheva and 4. Material and methods Smirnova, 2006; Middlemiss, 1981, 1984a, b; Lazar et al., 2010). The preparation of serial sections is an essential method for The material is preserved as articulated, but sometimes partly analyzing the internal shell structures of often externally very abraded, shells. To study the internal morphology, the preparation of similar specimens of Mesozoic brachiopod taxa. New digital serial sections was the only possible method. Computer tomographical reconstruction techniques allow detailed taxonomic identification scans have turned out to be useless because the composition of shell in spite of the strong homoplasy (see ‘Material and methods’). material and sediment matrix of these specimens does not allow Brachiopod genera of the family Sellithyrididae are geographi- sufficient differences for good images. However, the modern cally widely distributed and very abundant in Lower Cretaceous strata. Even though the majority of these taxa are externally very similar they represent good index fossils for this time span. The brachiopod genus Hadrosia is so far only reported from the Valanginian; as a result the occurrence of Hadrosia within the Rosablanca Formation confirms their Valanginian age. However, with the proposed Valanginian to ?early Hauterivian age of the Rosablanca Formation, the time span of this genus might extend also into the Hauterivian, but only when the precise biostratigraphy of the Rosablanca Formation in different places is known will it be possible to prove that the range of the genus in Colombia is restricted only to the Valanginian as has been indicated hitherto. Dietrich (1938) partly studied material of the fauna from Zapatoca, 200 km north of Santa Sofía, which contains a well-preserved assemblage dominated by Sellithyris sp., but the exact stratigraphical position of his material is uncertain and internal descriptions are not available. Sandy (1990,p.420,fig. 5.3aed) figured a specimen of Dietrich’s material which he assigned in a later work to the Saynoceras Horizon (cf. Sandy, 1991b,p.143).Thereareextensiveunexplored outcrops of shallow marine, Lower Cretaceous sediments in the eastern Andes of Colombia including Boyacá and Santander provinces and other similar sediments in Guajira Province. We assume that they are likely to containing a rich record of brachiopods including Fig. 3. Palaeogeographical map for the Early Cretaceous showing the occurrences of Hadrosia. It is necessary to state that even though no Sellithyris occurs taxa of Hadrosia and their migration way through the Tethys. (Map from Ron Blakey, in our material, it may be present in other localities in Colombia. NAU Geology; http://jan.ucc.edu). Author's personal copy

260 M. Schemm-Gregory et al. / Cretaceous Research 34 (2012) 257e267 laboratory techniques that we use to analyze the peels allow detailed 5. Systematic palaeontology 3D reconstructions of the internal features of the articulated specimens (e.g., Schemm-Gregory et al., 2008; Feldman et al., 2010; Phylum Brachiopoda Duméril, 1806 Schemm-Gregory, 2010). As a result, direct comparison of the Subphylum Rhynchonelliformea Williams et al., 1996 internal morphology of our specimens enables us to elucidate onto- Class Rhynchonellata Williams et al., 1996 genetical and phylogenetical relationships and, in a second step, to Order Terebratulida Waagen, 1883 contribute to new implications for palaeobiogeographical interpreta- Suborder Terebratulidina Waagen, 1883 tion. Several hundred serial sections of representative specimens were Superfamily Terebratuloidea Gray, 1840 prepared using a WOKO 50P grinding machine with slice-spacing of 50 Family Sellithyrididae Muir-Wood, 1965 and 100 mm. Acetate peels were used to record the morphological Subfamily Sellithyridinae Muir-Wood, 1965 information from each slice; these were subsequently digitized using a digital camera (Canon 300 D). Three-dimensional reconstruction methods are those of Sutton et al. (2001, 2005),whichwereimple- Remarks. In this study we consider the genera of the subfamily mented using the custom SPIERS software suite for registration, virtual Nerthebrochinae Cooper, 1983 to be members of the Sellithyridinae preparation, and interactive visualization. The digitized acetate peels Muir-Wood, 1965, because in our opinion the morphological were subsequently manually aligned with SPIERSalign and, in a second differences between the genera given in their diagnosis are not step, edited using different masks in SPIERSedit. The 3D images are sufficient for justify the erection of a subfamily (Cooper, 1983, p. 38, copied out of SPIERSview. SEM pictures were taken of uncoated Table 1); rather they represent differences at the genus level. specimens in a low pressure vacuum at 25 kbar and 15 kV and spot size Furthermore, the genus assignments to these two subfamilies, of 60 with a JSM-6490LV machine. Drawings of the peels were done Sellithyridinae and Nerthebrochinae, are interpreted differently in with help of a camera lucida. Measurements were taken with digital Cooper (1983, p. 38) and the revised “Treatise of Invertebrate callipers and rounded to 0.1 mm or using the free ImageJ 1.43u soft- Paleontology” (Lee and Smirnova, 2006, pp. 2062e2066). Subse- ware. Specimens were coated with ammonium chloride prior to quently, we propose the synonymization of the Nerthebrochinae. photography. The systematics follows the revised “Treatise on Inver- Additional studies are strongly recommended to prove our argu- tebrate Paleontology” (Lee and Smirnova, 2006). ment; however, the revision of the family Sellithyrididae is far beyond the scope of this work. Institutional abbreviations. UN-DG-BR, Departamento de Geociencias, Universidad Nacional de Colombia, Bogotá, Colombia; USNM, Genus Hadrosia Cooper, 1983 Smithsonian Institution/National Museum of Natural History, Washington DC, USA; YPM, Peabody Museum of Natural History, Type species. Hadrosia convexa Cooper, 1983, pp. 195, 196, pl. 17, figs. Yale University, New Haven, CT, USA. 36e41, pl. 67, figs. 8, 9.

Table 1 Morphological comparison of Hadrosia and Sellithyris.

Hadrosia gracilis sp. nov. Hadrosia convexa Cooper, 1983 Sellithyris Middlemiss, 1959 size small to medium medium to large small to medium outline elongate elliptical elongate elliptical pentagonal, almost as wide as long curvature equi- to gently equi- to gently usually ventribiconvex, rarely ventribiconvex ventribiconvex equibiconvex anterior margin strongly sulciplicate narrowly sulciplicate strongly sulciplicate foramen large, permesothyridid large, permesothyridid large, meso- to permesothyridid symphytium hardly concealed mostly concealed fully visible median sulcus short, deep short, deep highly variable median myophragm present unknown present lateral ridges present unknown absent/present loop angle 29e30 95e100 40e45 loop length and width 1/3 of length and width 1/3 of length and width slightly more than 1/3 of length and of dorsal valve of dorsal valve width of dorsal valve cardinal process small half ellipse small half ellipse half ellipse, wide, thin, indented medially socket ridges thin, erect, gently curved thin, erect, clearly arched thin, erect, gently curved sockets broad narrow broad fulcral plates small, without lateral unknown small, without lateral extensions extensions outer hinge plates wide, triangular, short, fairly wide, triangular, fairly wide, end in a point just below extend to the base of short, extend to the base (dorsal) of the crural process the crural process of the crural process crural bases thin, forming a high ridge broad, forming a high ridge elevated crural processes anterior to midloop, anterior to midloop, stoutly anterior to midloop, acutely pointed, sharply angular, directed angular, directed antermedially moderately long, venteromedially descending lamellae short, laterally curved, short, laterally curved, wide short, outwardly curved, narrow angle 30 transverse band broad, strongly arched, stout, broad, strongly arched, narrow, strongly arched, protuberant, steep lateral sloops, steep lateral slopes, 25 almost horizontal almost horizontal from horizontal median crest short, broadly rounded short, angular to rounded 1/3 of transverse band, narrow rounded terminal points extended, short and rounded extended, short and rounded not extended, rounded Geographic distribution Colombia Basses Alpes, France Great Britain, Central Europe, ?cosmopolitan Stratigraphic distribution Valanginian, ?Hauterivian Valanginian AptianeTuronian Author's personal copy

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Fig. 4. AeAF, Hadrosia gracilis sp. nov. AeI, UN-DG-BR 9449. A, ventral, B, dorsal, C, lateral, D, anterior and E, posterior views of articulated shell. F, oblique lateral view of loop. G, upper view of ventral internal shell. H, upper and I, oblique lateral views of dorsal internal shell. J-T, UN-DG-BR 9453. J, dorsal, K, ventral, L, anterior, M, lateral and N, posterior views of articulated shell. O, P, upper views of O, internal dorsal and P, ventral shell. Q, oblique posterior view of dorsal internal shell. R, oblique lateral view of loop. S, T, apical region of S, dorsal and T, ventral shell. Scale bars represent 2 mm in F, R, 5 mm in S, T. UeZ, UN-DG-BR 9455, holotype, in U, ventral, V, dorsal, W, anterior, X, Y, lateral and Z, posterior views of articulated shell. AAeAF, UN-DG-BR 9465, articulated shell in AA, AC, lateral, AB, anterior, AD, ventral, AE, posterior and AF, dorsal views. AGeAI, Hadrosia convexa Cooper, 1983. AG, AH, USNM 550930a, holotype in U, ventral and V, dorsal views. AI, USNM 550930e. Ventral view of loop on dorsal interior. All ﬁgures are 1.5 unless otherwise indicated. Author's personal copy

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Emended diagnosis. Medium-sized sulciplicate shells with a shallow comparison between Sellithyris and taxa of Hadrosia is given in umbonal chamber and an anterior rounded loop with a stout, broad Table 1. transverse band. (Modiﬁed after Cooper, 1983, p. 195). Hadrosia gracilis sp. nov. e Stratigraphic and geographic distribution. Valanginian, ?Hauterivian Figs. 4 7, Appendix (Lower Cretaceous); France and Colombia. 1985 Terebratulla sella Sowerby: Guzman, p. XII-5. Species included. Hadrosia convexa Cooper, 1983; Hadrosia gracilis v2009 Sellithyris sella (Sowerby): Rojas-Briceño and Patarrayo. sp. nov. v2010 Sellithyris cf. sella (Sowerby): Rojas-Briceño and Patarrayo, p. 91.

Remarks. Externally Hadrosia is most closely similar to Sellithyris Derivation of name. Latin, gracilis, thin, with reference to the elon- Middlemiss, 1959 within the Sellithyridinae Muir-Wood, 1965,but gate shell form. shows stronger convex valves and a more incurved beak. The loop in Hadrosia differs from Sellithyris in being at a narrower angle, and in Holotype. Articulated specimen illustrated in this work (Fig. 4UeZ) having a more anterior crural process, a more strongly transverse and housed in the Departamento de Geociencias, Universidad band that extends much closer under the crural process, and more Nacional de Colombia, Bogotá, Colombia, under the inventory number pronounced terminal points from the loop. A morphological UN-DG-BR 9455. Length 23.3 mm, width 19.2 mm, and thickness 12.5.

Fig. 5. SEM micrographs showing the calcitic ﬁbres of the secondary shell layer (AeE) and the punctae of the primary shell layer (F) of Hadrosia gracilis. A, B, UN-DG-BR 9449. CeF, UN-DG-BR 9453. Author's personal copy

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Fig. 6. Serial sections through Hadrosia gracilis.AeAC, UN-DG-BR 9453, all ﬁgures 3.0; sectioning perpendicular to commissural plane, sectioning distance in mm from posterior: bs, brachiopod shell; cb, crural base; cp, cardinal process; crpr, crural process; daf, dorsal adductor ﬁeld; dl, descending lamella; ds, dental socket; dv, dorsal valve; fp, fulcral plate; im, internal mould; m, myophragm; ohp, outer hinge plate; pf, pedicle foramen; pf, pedicle foramen; r, ridge; sr, socket ridge; t, tooth; tb, transverse band; uc, umbonal chamber; vv, ventral valve.

Type locality. Near “la Fábrica” cave and Las Juntas south of the town Stratigraphic and geographic distribution. As for genus. of Santa Sofía, Boyacá Province, central Colombia. Diagnosis. Elongated Hadrosia with a well-developed sulcus, a partly concealed symphytium, thin crural bases, a broadly Stratigraphic horizon. Upper Rosablanca Formation, Valanginian, rounded median crest, and a crural process directed perpendicular ?Hauterivian, Lower Cretaceous. to the commissural plane. Author's personal copy

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Fig. 7. Serial sections through Hadrosia gracilis.AeQ, UN-DG-BR 9449, all ﬁgures 3.0; sectioning perpendicular to commissural plane, sectioning distance in mm from posterior end: bs, brachiopod shell; cb, crural base; crpr, crural process; dl, descending lamella; ds, dental socket; dv, dorsal valve; fp, fulcral plate; im, internal mould; ohp, outer hinge plate; sr, socket ridge; t, tooth; tb, transverse band; tp, terminal point; vv, ventral valve.

Material. Eight articulated shells: UN-DG-BR 9449, 9453 (used for mid-length of shell. Fold within sulcus narrow, weak to strong, and serial sections), 9455 (holotype), 9460 (incomplete), 9465, 9468, clearly conspicuous. Shells swollen in umbonal regions of 9471,1001; three partly abraded external ventral valves: UN-DG-BR both valves, ventral beak erect. Pedicle foramen large and per- 9476, 1002, 1003. The material with measurements is given in mesothyrid. Pedicle collar clearly developed. Symphytium con- Appendix A. cealed by dorsal beak. Beak ridges not visible. SEM images obtained from non-coated specimens in a low Description. Shell: specimens small to medium-size, outline sub- vacuum mode at 15 kV show the primary layer (Fig. 5C) and the elliptical elongate in a longitudinal direction with maximum width calcitic ﬁbres of the secondary layer (Fig. 5). At the dorsal anterior at midlength or shortly anterior to it. Shells equibiconvex to margin well-preserved punctae across the calcitic ﬁbres show ventribiconvex in longitudinal section with smooth surface. a density of 197 punctae/mm2.(Fig. 5F). Anterior margin narrow sulciplicate at adult stage; in juvenile Interior of ventral valve: internal shell of ventral valve unspec- stages rectimarginate. Median sulcus short, begins anterior of tacular. Ventral sulcus and median fold clearly impressed. Small Author's personal copy

M. Schemm-Gregory et al. / Cretaceous Research 34 (2012) 257e267 265 and cyrtomatodont teeth are oriented at an angle of ca. 45 to each America and Western Europe in coincidence with the opening of other (Figs. 4T, 6R). the South Atlantic Ocean during the Early Cretaceous. Further Interior of dorsal valve: umbonal chamber shallow. Cardinal studies of that brachiopod bioevent including Hadrosia, Sellithyris, process small, outline inconspicuous in our material (Fig. 6K). and hitherto unidentified taxa in the Lower Cretaceous strata of the Dorsal diductor field comb-like striated (Fig. 6P). Socket ridges thin Andean region will provide and excellent opportunity for evalu- and gently curved over the broad dental sockets at the anterior end. ating the partially contradictory ideas about the climatic conditions Fulcral plates small, oriented parallel to commissural plane, and during ValanginianeHauterivian times (Weissert and Lini, 1991, without lateral extensions. Outer hinge plates thick, triangular, and 1998; Lini et al., 1992; Price, 1999; Aguirre-Urreta et al., 2008). In extending as a ridge in an interior direction towards the crural addition, an accurate taxonomic determination of brachiopod process of the deltiform and short loop, which is less than one third assemblages supported by high-resolution 3D reconstructions will of shell length. On their inside margin the thin crural bases form be useful to test the provinciality of low latitude Tethyan a high ridge almost perpendicular to the commissural plane and brachiopod faunas, which has been suggested by Middlemiss lead into the crural process. Crural processes sharp, pointing in (1984b). a ventromedian direction, and situated at or shortly anterior of mid-loop reaching into the ventral valve (Figs. 6Y, 7I, J). Descending Acknowledgements lamellae short, gently curved in a lateral direction and situated in an angle of 30 to each other. Transverse band broad, arched, and The research of MS-G in Panamá was partly financed by the with steep lateral slopes that are almost perpendicular to Smithsonian Tropical Research Institute. The visit of AR-B to the commissural plane (Figs. 4F, I, 7L, M). Terminal points short and Peabody Museum, Yale University was paid by a Charles Schuchert fi anterior rounded (Fig. 4F, R; Appendix B). Dorsal adductor eld and Carl O. Dunbar Grant. Both authors acknowledge the following at divided by a low, faint medium myophragm and one lateral ridge the Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt on either side of it (Fig. 6R, S). Measurements of specimens studied am Main, Germany: Bernhard Stribrny for providing access to the and of the loops are given in the appendices. laboratory; Tina Emmel and Michael Ricker for technical help; Claudia Franz for the SEM images. MS-G is grateful to Eberhard Discussion. The differences between the morphological features of Schindler of the same institute for providing her with working space. Hadrosia gracilis sp. nov. and the type species, Hadrosia convexa Michael Brett-Surman (USNM) took the photographs of the Cooper’s Cooper, 1983, justify the erection of a new species. Hadrosia gracilis material. PP thanks the company of geology students of the differs from H. convexa in smaller shells, a more conspicuous sulcus, Universidad Nacional de Colombia, during the field work in Santa a somewhat concealed symphytium, less curved socket ridges, Sofía. The manuscript has greatly benefited from the comments by broader sockets, wider outer hinge plates, a broadly rounded Robert B. Blodgett (Anchorage, USA) and Miguel O. Manceñido median crest, and a crural process that is oriented in a ventral (Museo Ciencias Naturales de La Plata, Argentina). direction. The type species is characterized by a mostly concealed symphytium, an angular to rounded median crest, and a crural References process that is oriented in an anteromedial direction. Furthermore, the descending lamellae are closer to each other in the new species Aguirre-Urreta, M.B., Price, G.D., Ruffell, A.H., Lazo, D.G., Kalin, R.M., Ogle, N., than in H. convexa. The transverse band in H. gracilis is almost Rawson, P.F., 2008. Southern Hemisphere Early Cretaceous (ValanginianeEarly horizontal, whereas it is inclined about 25 from horizontal in Barremian) carbon and oxygen isotope curves from the Neuquén Basin, Argentina. Cretaceous Research 29, 87e99. H. convexa. According to Pérez-Huerta et al. (2009), the density and Angiolini, L., Racheboeuf, P.R., Villarroerl, C.A., Concha, A.E., 2003. Stratigraphy and morphology of punctae is species-specific. At this stage of research brachiopod fauna of the Carboniferous El Imán Formation, Colombia. Revista Española de Paleontologia 18, 151e158. we could not study the shell material of H. convexa. Therefore, ’ fi d Archiac, J.A., 1847. Rapport sur les fossiles du Tourtia. In: Mémoires de la Société a further test for checking whether it is conspeci c (or not) may Géologique de France (2), 2, pp. 291e351. become available in the future when the shell fabric of the type Ballesteros, I., Nivia, A., 1985. La Formación Ritoque: registro sedimentario de una species of Hadrosia is studied in detail by SEM. A brachiopod that is albufera de comienzos del Cretácico. In: Etayo-Serna, F., Laverde, F. (Eds.), “ ” Proyecto Cretácico. Publicaciones Geológicas Especiales del Ingeominas 16, very similar externally to H. gracilis from the Neocomian of pp. XIV 1eXIV 17. Berklingen, Germany, is stored at the Peabody Museum of Natural Barret, S.F., 1988. The Devonian System in Colombia. In: McMillan, N.J., Embry, A.F., History, Yale University (YPB IP 227853). Without serial sections it Glass, D.J. (Eds.), Devonian of the World, Vol. 1. Regional Syntheses. Canadian Society of Petroleum Geologists, Calgary, Alberta, pp. 705e717. is impossible to say if it is a representative of Hadrosia. If it is, this Bogdanova, T.N., Hoedemaeker, Ph.J., 2004. Barremianeearly Albian Deshayesitidae, specimen would support our argument for the migration pathway Oppeliidae, Desmoceratidae and Silesitidae of Colombia. Scripta Geologica 128, of Hadrosia through the low latitude Tethys. 183e313. Bogotá, J., 1982. Contribución a la geología del oriente de los departamentos del A slightly older terebratulid “Terebratula” kanei Imlay, 1940 has Vichada y del Guainía (Colombia). Geología Norandina 6, 3e12. been reported from the Valanginian Barril Viejo shale of northern Bürgl, H., 1956. Catálogo de los amonitas de Colombia. Parte I. Pulchelliidae: Min- Mexico. It differs from Hadrosia externally in having a less isterio de Minas y Petrol, Instituto Geológico Nacional, República de Colombia. e conspicuous sulcus and a wider shell (Imlay, 1940). The internal Boletín de Geología IV (1), 1 119. Cadena, E., Gaffney, E., 2005. Notoemys zapatocaensis, a new side-necked turtle structures are unknown; as a result, a generic assignment accord- (Peurodira: Platychelyidae) from the Early Cretaceous of Colombia. In: ing to a modern palaeontological approach is not yet possible. This American Museum Novitates 3470, pp. 1e19. specimen might be a representative of Hadrosia or a related form, Cardozo, E.P., Ramirez, C.Q., 1985. Ambientes de depósito de la Formación Rosablanca: area de Villa de Leiva. In: Etayo-Serna, F., Laverde Montaño, F. depending on its internal morphology. (Eds.), Proyecto Cretácico. Publicaciones geológicas especiales del Ingeominas 16, pp. XIII 1e13. Caster, K.E., 1939. A Devonian fauna from Colombia. Bulletins of American e 6. Conclusions Paleontology 24 (83), 1 218. Catzigras, F., 1948. Sur une espèce hauterivienne de Térébratulideés suisse et méditerranéenne. Bulletin de la Société Géologique de France 18 Série 5, 18 New digital three-dimensional reconstruction techniques (6e7), 391e394. confirm the identification and presence of Hadrosia in South Cooper, G.A., 1983. The Terebratulacea (Brachiopoda), Triassic to Recent: a study of the brachidia (loops). Smithsonian Contributions to Paleobiology 50, 1e445. America. Its relationship to the Western European species supports Cooper, M.A., Addison, F.T., Alvarez, R., Coral, M., Graham, R.H., Hayward, A.B., the argument for a faunal pathway between northern South Howe, S., Martinez, J., Naar, J., Peñas, R., Pulham, A.J., Taborda, A., 1995. Basin Author's personal copy

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