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From the Late Devonian (Early Famennian) of Poland TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Zootaxa 3626 (3): 345–355 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3626.3.3 http://zoobank.org/urn:lsid:zoobank.org:pub:61A4F4E5-3F5A-4BA6-8535-DBF2A7E62226 A new Trimerocephalus species (Trilobita, Phacopidae) from the Late Devonian (Early Famennian) of Poland ADRIAN KIN1,2,4 & BŁAŻEJ BŁAŻEJOWSKI2,3 1Institute of Geological Sciences, Jagiellonian University, Oleandry 2a, 30-063 Cracow, Poland 2”Phacops” – Association of Friends of Geosciences, Targowa 29, Łódź 90-043, Poland. E-mail: [email protected] 3Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, 00-818 Warsaw, Poland 4Deceased Abstract This study presents a detailed morphological analysis of a new species belonging to the blind trilobite Trimerocephalus McCoy, 1849, T. chopini n. sp., based on exceptionally well preserved articulated specimens from the Late Devonian (Early Famennian) of the Holy Cross Mountains in central Poland. The occurrence of this taxon in Kowala Quarry near Kielce has been reported previously, with specimens often found in single-file queues representing migratory behaviour that was followed by a mass mortality event that preserved these assemblages. The new taxon is compared with other species of Trimerocephalus and is interpreted as being most closely related to a clade consisting of T. caecus, T. lelievrei, T. mimbi, T. shotoriensis and T. tardispinosus. Key words: Phacopinae, Late Devonian, Famennian, Holy Cross Mountains, Poland Introduction The blind phacopine genus Trimerocephalus McCoy, 1849 appears to have almost worldwide distribution in Late Devonian (Famennian) strata, except for America. Seventeen species of this genus are known (Crônier, 2003; Feist et al., 2009), six of which occur in Poland: T. caecus (Gürich, 1896), T. dianopsoides Osmólska, 1963, T. interruptus Berkowski, 1991, T. mastophthalmus (Richter, 1856), T. polonicus Osmólska, 1958, and T. (Trifoliops) trifolius (Osmólska, 1958). These species are widely distributed in the Fammenian carbonate strata of the Holy Cross Mountains (Osmólska, 1958, 1963; Radwański et al., 2009), while a single species is also found at Dzikowiec (formerly Ebersdorf) in Silesia (Lewowicki, 1959). Numerous specimens of Trimerocephalus were recently collected from yellowish marly shales of Early Famennian age that are exposed in the western part of the northern wall in Kowala Quarry (near Kielce in the Holy Cross Mountains, south-central Poland; Fig. 1). This occurrence was previously reported by Kin and Radwański (2008) and Radwański et al. (2009), but without detailed taxonomic and morphological analysis. The total thickness of the Famennian succession exposed in this quarry, which consists of five units (i.e., H–L = Palmatolepis triangularis to Palmatolepis expansa conodont zones), is approximately 180 m (see Berkowski, 2002). Age-diagnostic conodonts from the marly sediments—that also contain both monospecific trilobite queues and rare single specimens—are indicative of the lower part of the Palmatolepis marginifera Zone (see Radwański et al., 2009, p. 460; Unit I of Berkowski, 2002). The vast majority of Trimerocephalus specimens forming queues are partially or fully articulated and show considerable size variation (i.e., 5–20 mm in length). These trilobites are positioned on the tops of bedding surfaces, forming a single queue composed of a few to several specimens. The queue length is limited simply by the size of the joint-cracked stratum surface. The occurrence of the queues is restricted to two main horizons of marly shales, and a few queues were also found on the surface of some calcareous concretions, which occur irregularly within those horizons (Radwański et al., 2009). Successive horizons also contain isolated juvenile (Fig. Accepted by J. Paterson: 30 Jan. 2013; published: 14 Mar. 2013 345 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. 3A, C), sub-adult (Figs 2, 3B, D–F) and adult specimens (Fig. 4) that show identical morphologies with the sub- adult individuals represented in the queue assemblages (see Radwański et al., 2009). Specimens of Trimerocephalus from Kowala Quarry from were tentatively assigned to T. mastophthalmus by Radwański et al. (2009). Here we provide a much more detailed investigation of the morphology of this taxon and identify it as a new species, T. chopini n. sp. FIGURE 1. A: Map of Poland with the location of the Holy Cross Mountains region (grey rectangle). B: Generalised geologic map of the Holy Cross Mountains region and the location of Kowala Quarry (black arrow). C: Map of Kowala Quarry with the location of the studied section exposed in the western part of the northern quarry wall indicated by the black arrow. Material and methods The studied specimens (i.e., those preserved in queues and also single individuals, including the holotype) were collected by one of us (AK) in Kowala Quarry between 1995 and 2010. Most specimens are preserved as complete or nearly complete exoskeletons, representing both meraspid (Figs 2, 3) and holaspid stages (Fig. 4). Most show a variable degree of deformation. The collected material are housed at the Institute of Paleobiology, Polish Academy of Science in Warsaw (ZPAL Tr.8; old no. MGFA/Kow/TA). A total of around 450 specimens were studied and prepared manually at the Museum of Association of Friends of Geosciences, Łódź, including 78 trilobite queues. 347 individuals were subject to detailed measurements using vernier calipers with an accuracy of 0.01 mm and, in 346 · Zootaxa 3626 (3) © 2013 Magnolia Press KIN & BŁAŻEJOWSKI TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. part, using an advanced measurement technique associated with X-ray microcomputed tomography of the marls containing the trilobite queues. Scanning electron micrographs (e.g., Figs 2, 3, 4A–C) were taken using a Philips XL20 SEM at the Institute of Paleobiology, Polish Academy of Sciences (Warsaw, Poland). Specimens coated with ammonium chloride were photographed using a Canon EOS 400D digital camera (e.g., Fig. 4D, E). Morphological terminology follows Whittington et al. (1997), Crônier (2003) and Crônier et al. (2011). Ontogenetic nomenclature follows that of Crônier et al. (1998). Systematics Family Phacopidae Hawle & Corda, 1847 Trimerocephalus McCoy, 1849 Type species. Phacops mastophthalmus Richter, 1856. Trimerocephalus chopini n. sp. Figures 2–4 Material. Holotype, Tr.8/12.02.79, near complete exoskeleton (pygidium absent) of a late meraspid (M10) (Fig. 3F), from Unit I (late Early Famennian, Palmatolepis marginifera Zone) in Kowala Quarry, 7 km southwest of Kielce, Holy Cross Mountains, central Poland, 50° 47' 46.21" N 20° 33' 57.52" E. Etymology. In honour of Fryderyk Franciszek Chopin, the most famous Polish composer during the Romantic period of classical music, also called the ‘poet of the piano’. Diagnosis. Facial suture running along marginal furrow and not cutting cheek; the latter showing a very slight recess in contact with suture. Border furrows generally indistinct, but very shallow and wide near truncated genal angle. Ocular protuberance absent. Preoccipital furrow continuous along its length. Preoccipital ring straight, with two small, median tubercles. Occipital ring without median node. Pygidium trapezoidal with posterior margin almost rectilinear, width to length ratio of 2.8:1. Glabella and cheeks covered by very large to massive granules, usually uniform in size. Carapace covered by fine and moderately dense granulation. Description. Cephalon semicircular in outline, length to width ratio about 0.5. Glabella subpentagonal in outline, wide, rounded anteriorly with antero-lateral angles truncated (Figs 3F and 4A–C), length to width ratio about 0.6. Glabellar furrows S2 and S3 usually poorly developed, only visible on partially exposed internal moulds of glabellar area on three individuals [specimens: Kow / TA 76 (Radwański et al., 2009, pl. 2, fig. 3b) and Kow / TA 89, 90]; S2 short, bent obliquely upward and positioned very close to preoccipital furrow; posterior ramus of S3 very short and bent obliquely downward (as in T. caecus). Preoccipital furrow straight, well developed and of equal depth. Preoccipital ring straight, slightly convex with two small median tubercles. Occipital furrow slightly bent medially. Occipital ring wide, convex and without median node. Facial suture runs within the antero-lateral border furrow and does not extend onto the cheeks; cheeks exhibit a small recess in contact with suture (e.g., Figs 3F, 4A–C). In dorsal view, the margins of the glabella and cheeks are highly vaulted; cheek areas triangular in outline, with rounded to truncated posterolateral genal angles. Axial furrows deep and moderately wide, diverging forwards at an angle of about 55°. Lateral and posterior borders are widest at posterolateral corner of cephalon. Vincular furrow in holaspid moderately deep, straight medially, strongly directed transversely with smooth edges (Fig. 4D–E). In the genal part it is directed posteriorly (parallel to the cheek), and is very slightly crenulated with small, shallow pits (Fig.
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