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Hoplolichoides, Allolichas, Autoloxolichas

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Hoplolichoides, Allolichas, Autoloxolichas Geobios 35 (2002) 111–125 www.elsevier.com/locate/geobios Hoplolichoides, Allolichas, Autoloxolichas and Akantharges, and the classification of lichid trilobites Hoplolichoides, Allolichas, Autoloxolichas et Akantharges, et la classification des trilobites Lichidae David J. Holloway a,*, Alan T. Thomas b aInvertebrate Palaeontology, Museum Victoria, PO Box 666E, Melbourne, Victoria 3001, Australia bSchool of Earth Sciences, University of Birmingham, Edgbaston, Birmingham, B15 2TT, United Kingdom Received 9 November 2000; accepted 24 July 2001 Abstract Hoplolichoides PHLEGER, 1936 and Autoloxolichas PHLEGER, 1936 are revised based on redescription of their type species, H. conicotuberculatus (NIESZKOWSKI, 1859) and A. sanctamathiae (SCHMIDT, 1885) respectively. Hoplolichoides is considered to be most closely related to Hoplolichas DAMES, 1877, and both are included in a revised Homolichinae. Autoloxolichas is restricted to its type species, and other species previously assigned to the genus are placed in Allolichas KRUEGER, 1992, the concept of which is expanded. Autoloxolichas and Allolichas are included together with three other genera and subgenera in the Subfamily Platylichinae, previously considered to be a junior synonym of Homolichinae. The hypostomal morphology previously regarded as important in uniting homolichine and platylichine genera is now regarded as primitive for the Lichidae. Metopolichas GÜRICH, 1901, previously included tentatively in the Homolichinae on the basis of its hypostomal morphology, is reassigned tentatively to the Lichinae. The trochurine ‘Acanthopyge’ erbeni MEISCHNER, 1965, from the Givetian of Germany, is assigned to Akantharges PHLEGER, 1936, permitting clarification of the pygidial morphology of the genus. Akantharges and Ceratarges GÜRICH, 1901 are unique amongst lichids in having a curved, transverse ridge on the posterolateral cranidial lobe, suggesting that these genera may have been derived from a common ancestor. © 2002 Éditions scientifiques et médicales Elsevier SAS. All rights reserved. Résumé Les genres Hoplolichoides PHLEGER, 1936 et Autoloxolichas PHLEGER, 1936 sont révisés sur la base d’une redescription de leurs espèces types respectives: H. conicotuberculatus (NIESZKOWSKI, 1859) et A. sanctamathiae (SCHMIDT, 1885). Hoplolichoides est considéré comme le genre le plus proche de Hoplolichas DAMES, 1877. Les deux genres sont inclus dans les Homolichinae également révisés. Autoloxolichas ne contient que son espèce-type et d’autres espèces rapportées aurapavant à Autoloxolichas sont placées dans Allolichas KRUEGER, 1992 dont la définition est ainsi étendue. Autoloxolichas et Allolichas sont inclus avec trois autres genres et sous-genres dans la sous-famille des Platylichinae que l’on considérait avant comme un synonyme des Homolichinae. La morphologie de l’hypostome considérée autrefois comme importante (unissant les genres homolichines et platylichines) est à présent considérée comme primitive pour les Lichinae. Metopolichas GÜRICH, 1901 qui avait été placé dans les Homolichinae sur la base de sa morphologie hypostomale est placé dans les Lichinae. Le trochuriné ‘Acanthopyge’ erbeni MEISCHNER, 1965 du Givétien de l’Allemagne est attribué à Akantharges PHLEGER, 1936 permettant une classification de la morphologie pygidiale du genre. Akantharges et Ceratarges GÜRICH, * Corresponding author. E-mail addresses: [email protected] (D.J. Holloway), [email protected] (A.T. Thomas). © 2002 Éditions scientifiques et médicales Elsevier SAS. All rights reserved. PII:S0016-6995(02)00014-1 112 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125 1901 sont uniques chez les lichidés par leur crête transversale courbe sur le lobe cranidial postéro-latéral, suggérant que ces deux genres dérivent d’un ancêtre commun. © 2002 E´ ditions scientifiques et médicales Elsevier SAS. Tous droits réservés. Keywords: Trilobites; Lichidae; Systematics; Ordovician; Devonian Mots clés: Trilobites; Lichidae; Systématique; Ordovicien; Dévonien 1. Introduction species of Akantharges from the Devonian of Morocco was illustrated by Thomas and Holloway (1988, pl. 14, figs 299, The trilobite genera Hoplolichoides, Autoloxolichas and 200, 303), who based their concept of the genus largely on Akantharges, all erected by Phleger (1936), were reviewed that specimen. We have subsequently been able to examine by Thomas and Holloway (1988) in their revision of the the material of ‘Acanthopyge’ erbeni MEISCHNER, 1965 Lichida. Additional information on these genera has become from the Givetian of Germany, and recognised that this available subsequently, providing the opportunity to clarify species belongs to Akantharges. The specimens of erbeni their taxonomic concepts as well as to assess their likely are relatively well preserved and clarify some of the relationships. In the light of our new data, we assign characters of the genus, particularly pygidial characters. Hoplolichoides to a revised Homolichinae. The Subfamily Platylichinae is resurrected to accommodate Allolichas and Autoloxolichas, together with other genera. 2. Systematic palaeontology Hoplolichoides has not generally been accepted as a distinct genus, most authors (Warburg, 1939; Hupé, 1953; Morphological terminology used here is that of Thomas Tripp, 1957, 1959; Balashova, 1960) considering it to be a and Holloway (1988). Stratigraphical nomenclature follows junior subjective synonym of Hoplolichas DAMES, 1877. the revised Ordovician series proposed by Fortey et al. Thomas and Holloway (1988) recognised Hoplolichoides (1995, 2000). but expressed reservations about its utility as an indepen- Several of the species dealt with herein were documented dent genus, stating that the pygidial morphology was by Schmidt (1885). Until recently, the specimens from this insufficiently known for a full comparison with other and other parts of Schmidt’s series of trilobite monographs genera. In the present paper, we redescribe cephala and were not available for study. Photographs of most of pygidia of the type species, H. conicotuberculatus (NIESZ- Schmidt’s specimens are now available on the World Wide KOWSKI, 1859), and compare it with other species con- Web at the site entitled ‘Catalogue of the trilobites figured in sidered to be congeneric, showing that Hoplolichoides Friedrich Schmidt’s ‘Revision der ostbaltischen silurischen warrants recognition as a separate genus. Trilobiten’ (1881–1907)’ by D.L. Bruton, O.A. Hoel, Autoloxolichas was based on Lichas sanctamathiae L.T. Beyene and A. Yu. Ivantsov (internet address SCHMIDT, 1885, and that was the only species listed by http://www.toyen.uio.no/palmus/schmidt/index.html). The Phleger (1937) as belonging to the genus. The genus was specimens are also listed but not illustrated in Bruton et al. briefly diagnosed by Phleger (1936) as differing from (1997). Platylichas in lacking the continuation of the longitudinal Repositories of specimens are denoted by the following furrow behind the bullar lobe, but Öpik (1937) dismissed abbreviations: CNIGR, Central Scientific-Research Geo- this supposed difference as being due to a misunderstanding logical Exploration Museum named after F.N. Chernyshev, and placed sanctamathiae in Platylichas. Tripp (1957, St Petersburg; GIE, Institute of Geology, Tallinn Technical 1959) and Dean (1974) also considered Autoloxolichas to be University; Gö, Geologisch-Paläontologishes Institut und a junior synonym of Platylichas. Thomas and Holloway Museum, Georg-August-Universität, Göttingen; MB, Mu- (1988) recognised Autoloxolichas as a distinct genus to seum für Naturkunde, Humboldt Universität, Berlin; RM, which they assigned, in addition to sanctamathiae, a num- Naturhistoriska Riksmuseet, Stockholm. ber of species resembling ‘Lichas’ laxatus MCCOY, 1846. Family: LICHIDAE Hawle & Corda, 1847 We now believe, based on the study of plaster replicas of the Subfamily: HOMOLICHINAE Phleger, 1936 type material of sanctamathiae, that Autoloxolichas should Diagnosis – S1 effaced or almost effaced, so that bullar be restricted to that species, and that the other species lobe is more or less completely fused with L1b; resulting included in the genus by Thomas and Holloway (1988) composite lobe circumscribed by furrows, except in Leioli- should be placed in Allolichas KRUEGER, 1992, the chas which has longitudinal furrow effaced externally. concept of which should be expanded. Palpebral lobe narrow, of almost uniform width throughout, Akantharges was until relatively recently known only moderately curved in outline. Hypostome with middle body from the type species, Lichas gourdoni BARROIS, 1886, as wide across posterior lobe as across anterior lobe, or the material of which is strongly deformed and difficult to wider; shoulder situated level with posterior border furrow; interpret. A well preserved cranidium of an undescribed border greatly expanded and flap-like at and behind D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125 113 shoulder. Pygidium with two to three pairs of pleural furrows and six to eight marginal spines (spines absent in Leiolichas); anterior and posterior pleural bands flattened. Genera included – Homolichas SCHMIDT, 1885; Conolichas DAMES, 1877; Hoplolichas DAMES, 1877; Hoplolichoides PHLEGER, 1936; Leiolichas SCHMIDT, 1885; Otarozoum THOMAS & HOLLOWAY, 1988. Remarks – S1 is effaced, or almost so, in Conolichas, Hoplolichas, Hoplolichoides and Otarozoum, the bullar lobe almost completely fusing with L1b. The resulting composite lobe is circumscribed by the longitudinal and axial
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