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Devonian trilobite Cordania from Australia, The Ebach, Malte C ABSTRACT-The genus Cordania Clarke, 1892, has been known from the Lower Devonian (Lochkovian-Pragian) of the Appalachian Province and apparently from the Lower-Middle Devonian of China. Probable Lochkovian strata of the Biddabirra Formation in the Amphitheatre Group from near Cobar, New South Wales, Australia, have yielded Cordania buicki new species, extending the range of the genus to eastern Australia. Cladistic analysis of Cordania identifies C. buicki as more closely related to species from Oklahoma than to a northern Appalachian grade. THE BRACHYMETOPID trilobite Cordania has been studied in detail by Whittington (1960), Campbell (1977), and Adrain and Kloc (1997). It includes at least eight species, seven of which are from the Appalachian Province between Quebec and Oklahoma, as well as two probable congenerics from China (Zhou, 1989). Cordania buicki n. sp. now extends the distribution of Cordania to Australia. LOCALITY AND GEOLOGICAL CONTEXT The new species of Cordania is part of a fauna including twelve trilobite species from a locality called The Bluff, 60 km southwest of Cobar, western New South Wales (Fig. 1). The site is located 1.5 km southwest of The Bluff homestead (locality F5 of Glen, 1987, grid reference GR 713815). The trilobites occur in the Biddabirra Formation of the Amphitheatre Group in the Cobar Supergroup. In the study area, the Biddabirra Formation has a thickness of about 1.5 km, and lies stratigraphically between the lower Amphitheatre Group and the upper Amphitheatre Group (Glen, 1987). It consists of poorly outcropping, medium to thick bedded sandstones. Trilobites are preserved as molds, mostly disarticulated, in medium grained sandstone. Petrographically, Glen (1987) described this lithology in the Biddabirra Formation as a (sub)lithic arenite, and attributed its characteristic facies to deposition from proximal turbidites. The presence of the brachiopod Howellella jaqueti in the Biddabirra Formation was regarded as evidence for a Pragian age, possibly extending down into the Lochkovian (Sherwin cited in Glen, 1987). A more recent assessment of brachiopods in the Cobar Supergroup (Sherwin, 1995) recognizes the occurrence of H. jaqueti as probably Lochkovian. Trilobites associated with Cordania buicki n. sp. in the Biddabirra Formation, currently under study by Ebach, include Kainops cf. ekphymus (Jones et al., 1986), new species of Paciphacops, Leonaspis, and Acanthopyge (Lobopyge), species of Crotalocephalus, Scotoharpes, Cornuproetus, Gerastos, and Cyphaspis, a tropidocoryphid, and a styginid. Although a Pragian age cannot be ruled out based on these taxa, close comparisons can be made with species recognized as Lochkovian in other parts of New South Wales (e.g., Kainops ekphymus, from the lower part of the Tangerang Formation in the Windellama district; Jones et al.,1986). OCCURRENCE OF CORDANIA OUTSIDE NORTH AMERICA AND AUSTRALIA Zhou (1989) reassigned two Chinese species to Cordania. Of these, C. transversus Nan, 1976 (type species of Latecephalus Nan, 1976 but synonymized by Zhou), occurs in Inner Mongolia, and the other, C. junggarensis (Xiang and Zhang, 1983), occurs in western Junggar, Xinjiang. The occurrence of Chinese species would appear to shed light on a distribution pattern otherwise restricted to eastern North America and Australia. Although illustrated material of C. transversus (Nan, 1976, pl. 201, figs. 5, 6) also resembles Mystrocephala (Adrain and Kloc, 1997), we tentatively accept Zhou's (1989) referral of it to Cordania. The species has deep epiborder and anterior border furrows, more typical for Cordania, while pygidial rib structure (equal height of the anterior and posterior pleural bands) more closely resembles that of Cordania than Mystrocephala (posterior bands prominently raised distally). A lateral view of the cranidium is required to code the species' characters and ascertain its synonymy, hence it has not been included in the phylogenetic analysis. Alberti's (1969) description of a pygidium of Cordania from the Lower Devonian of Morocco was discussed and rejected by Campbell (1977), who considered the specimen to possess no distinctive features that justify placement in Cordania or, indeed, the Cordaniinae. PHYLOGENETIC ANALYSIS OF CORDANIA Data.-Eleven characters used for discerning relationships between seven species of Cordania are listed in the Appendix. The sources consulted for coding are Whittington (1960), Campbell (1977), and Adrain and Kloc (1997). The character matrix used in the analysis is shown in Table 1. The data were run on PAUP version 3.1.1 (Swofford, 1993) on a Power Macintosh, using the Branch and Bound search option with "furthest" addition and ACCTRAN (accelerated transformation) optimization. Multistate characters were treated as unordered, and multistate taxa as polymorphic. Mystrocephala Whittington, 1960, was used as outgroup due to its close resemblance to, and inferred relationship with, Cordania. Campbell's (1977) classification implies a closer relationship between Cordania and Radnoria Owens and Thomas, 1975, based largely on the form of the posterior pygidial pleural ribs. Campbell (1977, p. 22) allowed that Mystrocephala could be derived from a cordaniine (i.e., Cordaniinae is paraphyletic) and a subfamily composed of Radnoria and Cordania has not been employed in recent studies. However, to test the sensitivity of

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our results to alternative outgroups, we also analyzed the data with Radnoria as outgroup, coding based on R. syrphetodes Owens and Thomas, 1975. Results.-Rooting on Mystrocephala yielded a single most parsimonious cladogram of 18 steps with a consistency index (CI) of 0.889 and a retention index (RI) of 0.846 (Fig. 2). The alternative rooting on Radnoria produced the same topology for species of Cordania, of equal length, with a slightly lower retention index (0.818). The Australian Cordania buicki n. sp. is nested within North American clades, being resolved as most closely related to C. wessmani and C. falcata. As formalized by Adrain and Kloc (1997), these latter two species show enough variation to justify separating Campbell's (1977) two types (A and B) into two species. They are, however, resolved as each other's closest relative, as predicted by Adrain and Kloc (1997). Given the low number of characters, branch support (Bremer, 1994) is not strong. Only the node uniting all taxa except C. cyclurus is present in all (ten) trees one step longer than the shortest, and this node is collapsed in the strict consensus of all (37) trees two steps longer than the shortest. Biogeography.-The cladogram (Fig. 2) suggests that the northern part of the Appalachian Province is the most likely ancestral area for Cordania. Its probability can be expressed using Bremer's (1992) ancestral area method. This method is based on comparing the numbers of necessary gains of an area on the cladogram if that area is not part of the ancestral distribution versus the number of losses of the area if it were part of the ancestral distribution. The gain/loss (GIL) ratios for the northern Appalachian region, southern Appalachian region, and Australia are 4, 0.2, and 0.2, respectively. The rescaled gain/loss quotients (AA of Bremer, 1992) are 1, 0.05, and 0.05, respectively. The northern part of the Appalachian Province can thus effectively be regarded as part of the ancestral area, whereas this is much less probable for Australia and Oklahoma. Presence of Cordania in these areas may be attributed to dispersal. Trilobite species associated with Cordania buicki in the Biddabirra Formation mostly have close relatives in other eastern Australian Lochkovian-Pragian trilobite faunas (e.g., Chatterton et al., 1979; Holloway and Neil, 1982). Cordania is distinctive in indicating affinities to the Kazakhstan-Appalachian province of Zhou (1989). SYSTEMATIC PALEONTOLOGY Specimens figured in this paper are housed in the type collections of the Palaeontology Section, Australian Museum, Sydney (prefixed AMF). Terminology.-We follow the interpretation of Campbell (1977) and Adrain and Kloc (1997) that a furrow on the outer part of the cephalon in Cordania and other brachymetopids is an epiborder furrow, impressed upon the cephalic border. The terms "anterior area of anterior border" and "posterior area of anterior border" are used in this work to refer to parts of the anterior cranidial border in front of and behind the epiborder furrow, respectively. FamilY BRACHYMETOPIDAE Prantl and Pribyl, 1951 Discussion.-Adrain and Kloc (1997) have summarized competing views on subfamilial classifications within the Brachymetopidae (Owens and Thomas, 1975; Campbell, 1977; Owens and Hammann, 1990). As currently understood (Adrain and Chatterton, 1993), Brachymetopidae includes the following nine genera: Brachymetopus McCoy, 1847; Cordania Clarke, 1892; Cheiropyge Diener, 1897; Tschernyshewiella Toll, 1899; Proetides Walter, 1924; Australosutura Amos, Campbell and Goldring, 1960; Mystrocephala Whittington, 1960; Radnoria Owens and Thomas, 1975; Loeipyge Kobayashi and Hamada, 1979. Genus CORDANIA Clarke, 1892 Type species.-Phaethonides cyclurus Hall and Clarke, 1888, from the New Scotland Limestone (Lochkovian), Lower Helderberg Group, New York, by original designation; revised by Whittington (1960). Diagnosis.-See Campbell (1977). Occurrence.-Cordania occurs in the Lochkovian and Pragian in the Appalachian Province between Quebec and Oklahoma (Whittington, 1960; Campbell, 1977; Adrain and Kloc, 1997), in the Lochkovian in western New South Wales, Australia, and probably in Xinjiang, China, and Inner Mongolia (Zhou, 1989). CORDANIA BUICKI new species Figure 3 Diagnosis.-Cordania buicki is distinguished from all congenerics by the steep slope, distinct convexity, and strong roll of the long (sag.) anterior cranidial border; anterior area of anterior border nearly vertical; epiborder furrow narrow, well-impressed. Description.-Glabella with subangular anterolateral corners; L1 distinct, triangular, with subangular corners, convex (tr.) and half as high as the glabella; St moderately deep near axial furrow, abruptly shallowing posteromedially, completely isolating L1 (Fig. 3.2). Glabella strongly convex (sag.), but only curving on frontal lobe and towards SO; mid surface weakly convex and sloping at approximately 45 degrees. S2 a narrow (tr.) notch. L2 short (exsag.), weakly inflated. Shallow anteromedially directed S3 visible on some internal molds. Axial furrow narrow and moderately deep against LI, distinctly widened anterior to S1, with small fossula anteriorly. SO short (sag., exsag.), shallow. Occipital ring longest medially; tubercles evenly scattered over occipital ring, with largest tubercles medially. Preglabellar furrow of moderate length and

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depth, with numerous large, shallow, linear pits (Fig. 3.1). Anterior border furrow of variable depth, from shallow but distinct (Fig. 3.1) to moderately deep (Fig. 3.7), merged with preglabellar furrow medially, without preglabellar field. Epiborder furrow narrow, sharply impressed, separating anterior border into two unequally sized parts, forming two distinct convex rises. Border evenly lengthening medially. On holotype (Fig. 3.2), frontal area 30 percent length (sag.) of cranidium; sagittal length of cranidium 70 percent width across anterior border. Palpebral area steep, higher than frontal area of fixigena. Eye ridge faint. Postocular fixigena steeply sloping to posterior border furrow. Anterior section of facial suture a distinct 'S' shaped curve, running posteriorly from alpha, crossing epiborder furrow at beta, then straight and running nearly adaxially before sharply curving back to gamma; from gamma the facial suture runs posteriorly to palpebral lobe, moderately convex outward at delta; posterior section of facial suture runs posterolaterally from epsilon to the posterior border. Tubercles scattered over median lobe of glabella, largest centrally; a few tubercles on LI; glabellar tubercles smaller than those on border; large tubercles in distinct rows along posterior area of anterior border (Fig. 3.1, 3.2); tubercles slightly smaller on anterior area of border, absent on anteroventral part. Small pits distinct and dense inside epiborder furrow, present but less prominent on anterior area, becoming indistinct toward anterior cranidial margin, preglabellar furrow and palpebral lobe. Length of genal spine about 150 percent width (tr.) of librigenal field and border. Large tubercles evenly scattered over librigena in same manner as on the cranidium. Internal mold shows wide band of numerous terrace lines along doublure. Both lateral border and doublure are convex and wide; doublure flattened beneath genal spine. Lateral border gently narrowing towards end of genal spine. Hypostome and rostral plate unknown. Pygidium semicircular in outline, length about 50 percent of width. Axis parabolically arched (tr.), raised high above pleural field, gently convex (sag.) anteriorly, with convexity increasing behind fifth ring; posterior region steeply sloping. Eight or nine rings defined by clearly impressed ring furrows, two more weakly defined in terminal piece; five medially continuous ring furrows with slot-like incision distally. Axial furrow narrow, very shallow against posterior rings, indistinct behind axial terminus. Eight pleural ribs. Pleural furrow slightly longer (exsag.) than interpleural furrow, both sharply impressed. Anterior pleural band longer than posterior band distally; posterior band slightly more elevated. Pleurae moderately turned down distal to fulcrum. Pleural ribs become faint and turned inwards (tr.) around terminal piece, with last two ribs running almost straight back (exsag.). Postaxial region concave. Border narrow, without border furrow. Pleurae with small, subdued tubercles; axis with row of weak tubercles; faint median nodes on some axial rings. Etymology.-After Dr. Roger Buick. Types.-Holotype cranidium AMF 100287 (Fig. 3.2-3.4); figured paratypes AMF 100280-100286. All type material from the Biddabirra Formation (Lochkovian), The Bluff, 60 km southwest of Cobar, New South Wales, Australia. Other material examined.-65 cranidia, 54 pygidia, two poorly preserved thoracic segments, and 13 librigenae, none articulated to cranidia. All from type locality. Occurrence.-Biddabirra Formation, type locality. Discussion.-As noted in the diagnosis, no other species of Cordania possesses such a steep anterior area of the anterior border as C. buicki. Compared to C. falcata Whittington, 1960, and C. wessmani Adrain and Kloc, 1997, C. buicki has shorter genal spines, a more steeply sloping glabella, and a more elevated pygidium. It is further distinguished from C. wessmani by its marked border roll, shallower anterior cranidial border furrow, and much deeper epiborder furrow. Cordania buicki differs from C. macrobius (Billings, 1869) (see Whittington, 1960, for revision) in having larger tubercles, stronger elevation of the glabella, and a more sharply impressed but shorter (sag., exsag.) epiborder furrow. Cordania buicki displays a far longer posterior area of the anterior border than C. becraftensis Clarke, 1900, and C. gasepiou Clarke, 1908 (both revised by Whittington, 1960), and a steep (rather than rimlike) anterior area. The type species, C. cyclurus (Hall and Clarke, 1888), displays a deeper occipital furrow and S 1 than C. buicki, has a depressed, strongly pitted posterior area of the anterior border (this depressed region apparently incorporating the epiborder furrow), and bears more prominent pygidial tubercles. An unnamed species of Cordania from Tennessee (Whittington, 1960, pl. 52, fig. 4) has a much shorter posterior area of the anterior border than that of C. buicki. ACKNOWLEDGMENTS We wish to thank C. and J. Osgood for letting us collect on their property, and the Australian Museum for providing a Postgraduate Research Fellowship to M.C.E. for funding of a field trip. The Journal's referees provided useful suggestions. We are deeply grateful for the help of R. Buick. REFERENCES ADRAIN, J. M. AND B. D. E. CHATTERTON. 1993. A new rorringtoniid trilobite from the Ludlow of Arctic Canada. Canadian Journal of Earth Sciences, 30:1634-1643. -, AND G. J. KLoc. 1997. Lower Devonian aulacopleuroidean trilobites from Oklahoma. Journal of Paleontology, 71:703-712. ALBERTI, G. K. B. 1969. Trilobiten des jungeren Siluriums sowie des Unter- und Mitteldevons. I. Mit Beitrigen zur Silur-Devon-Stratigraphie einiger Gebiete Marokkos und oberfrankens. Senckenbergishen Naturforschenden Gesellschaft, 520, 692 p. AMOS, A. J., K. S. W. CAMPBELL, AND R. GOLDRING. 1960. Australosutura gen. nov. (Trilobita) from the Carboniferous of Australia and

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Argentina. Palaeontology, 3:227-236. BILLINGS, E. 1869. Description of some new species of fossils with remarks on others already known, from the Silurian and Devonian rocks of Maine. Proceedings of the Portland Society of Natural History, 1:104-126. BREMER, K. 1992. Ancestral areas: a cladistic reinterpretation of the center of origin concept. Systematic Biology, 41:436-445. -. 1994. Branch support and tree stability. Cladistics, 10:295-304. CAMPBELL, K. S. W. 1977. Trilobites of the Haragan, Bois d'Arc and Frisco Formations (Early Devonian) Arbuckle Mountains Region, Oklahoma. Oklahoma Geological Survey Bulletin, 123:1-227. CHATTERTON, B. D. E., B. D. JOHNSON, AND K. S. W. CAMPBELL. 1979. Silicified Lower Devonian trilobites from New South Wales. Palaeontology, 22:799-837. CLARKE, J. M. 1892. On Cordania, a proposed new genus of trilobites. New York State Museum, 45th Annual Report for 1891:440-443. 1900. The Oriskany fauna of Becraft Mountian, Columbia County, N.Y. New York State Museum Memoir, 3, 128 p. 1908. Early Devonian history of New York and eastern North America. New York State Museum Memoir, 9, 366 p. DIENER, C. 1897. The Permocarboniferous of Chitichun No. 1. Memoirs of the Geological Survey of India. Palaeontologica Indica, Series 15, Himalayan Fossils, 1:1-105 GLEN, R. A. 1987. Geology of the Wrightville 1:100,000 sheet 8034. New South Wales Geological Survey, Sydney. HALL, J., AND J. M. CLARKE. 1888. Trilobites and other Crustacea. Natural History of New York, Paleontology, Volume 7, supplement to 5:236-277. HOLLOWAY, D.J., AND J. V. NEIL. 1982. Trilobites from the Mount Ida Formation (Late Silurian-Early Devonian), Victoria. Proceedings of the Royal Society of Victoria, 94:133-154. JONES, B. G., C. G. HALL, A. J. WRIGHT, AND P E CARR. 1986. The geology of the Bungonia-Windellama area, New South Wales. Proceedings of the Linnean Society of New South Wales, 108:267-286. KOBAYASHI, T., AND T. HAMADA. 1979. Permo-Carboniferous trilobites from Thailand and Malaysia. Geology and Palaeontology of SE Asia, 20:1-21. McCoY, E 1847. On the fossil botany and zoology of the rocks associated with the coal of Australia. Annals and Magazine of Natural History, 20:145-57, 226-236, 298-312. NAN, R.-S. 1976. Trilobita, p. 333-352. In Bureau of Geology, Inner Mongolia and North East China Institute of Geology (eds.), Paleontological Atlas of North China. Inner Mongolia, Volume 1, Paleozoic. Geological Publishing House, Beijing. OWENS, R. M., AND A. T THOMAS. 1975. Radnoria, a new Silurian proetacean trilobite, and the origins of the Brachymetopidae. Palaeontology, 18:809-822. -, AND W. HAMMANN. 1990. Proetide trilobites from the Cystoid Limestone (Ashgill) of NW Spain, and the suprageneric classification of related forms. Palaontologische Zeitschrift, 64:221-244. PRANTL, E, AND A. PRIBYL. 1951. A revision of the Bohemian representatives of the family Otarionidae R. and E. Richter (Trilobitae). Sbornik Statniho Geologickeho ustavu Ceskoslovenske Republiky, 17:353-512. SHERWIN, L. 1995. Siluro-Devonian brachiopods from the Amphitheatre Group (Cobar Supergroup), western New South Wales. Association of Australasian Palaeontologists, Memoir 18:61-96. SWOFFORD, D. L. 1993. PAUP: Phylogenetic Analysis Using Parsimony, Version 3.1. Program distributed by the Illinois Natural History Survey, Champaign. TOLL, E. VON. 1899. Beitrage zu kenntnis des Sibirischen Cambrian. Zapiski Imperatorskoi Akademii Nauk po Fiziko-Matematischeskomu Otdeleniyu, St. Petersburg, 8:1-57 WALTER, O. T 1924. Trilobites of Iowa and some related Paleozoic forms. Iowa Geological Survey Annual Reports 1923 and 1924, 31: 167-400. WHITTINGTON, H. B. 1960. Cordania and other trilobites from the Lower and Middle Devonian. Journal of Paleontology, 34:405-420. XIANG, L.-W., AND ZHANG, T-R. 1983. Trilobita, p. 534-556, 769-772. In Institute of Geological Sciences, Xinjiang Bureau of Geology (eds.), Palaeontological Atlas of Northwest China, Xinjiang Autonomous Region, Volume 2. Geological Publishing House, Beijing. ZHOU, Z.-Q. 1989. Occurrence of Devonian trilobite Cordania Clarke, and its palaeozoogeographical significance. Acta Palaeontologica Sinica, 28:550-552. ACCEPTED 17 NOVEMBER 1998 MALTE C. EBACH1,2 AND GREGORY D. EDGECOMBE1 1 Australian Museum, 6 College Street, Sydney South, New South Wales 2000 Australia, and Department of Geology and Geophysics, University of Sydney, New South Wales 2006, Copyright Paleontological Society May 1999 Provided by ProQuest Information and Learning Company. All rights Reserved

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