[Palaeontology, Vol. 49, Part 1, 2006, pp. 113–132]

THE FISH GENERA STREPSODUS AND ARCHICHTHYS (: RHIZODONTIDA): CLARIFYING 150 YEARS OF CONFUSION by JONATHAN E. JEFFERY Institute of Biology, University of Leiden, Postbox 9516, 2311GP Leiden, The Netherlands; e-mail: [email protected]

Typescript received 27 August 2003; accepted in revised form 13 December 2004

Abstract: The Carboniferous genus Strepsodus contains Nova Scotia, Canada) and more recently in Australia several described species, but has been mired in taxonomic (Ducabrook, Queensland). Rhizodontid remains from two confusion for nearly 150 years. The little-known genus Ar- Upper sites (one in Colombia, one in Turkey) chichthys (with two described species) is usually treated as have been attributed to Strepsodus, due to a misunder- a junior synonym of Strepsodus. These problems have been standing of the defining characteristics (autapomorphies) caused by, among other things, the poor quality of most of the genus. This paper reviews what is known of Strepso- specimens, and a heavy reliance on tooth and scale mor- dus and Archichthys, and advocates that each be treated as phology when erecting new species. Both genera are a monospecific genus until reliable morphological evidence known from Upper and Lower Carboniferous river and of further speciation is found. A neotype specimen for lake deposits in the UK, particularly coal shales, cement- Strepsodus sauroides is proposed, and a check-list of pub- stones and freshwater limestones. However, Strepsodus has lished rhizodontid species is appended. also been discovered at a number of North American localities (e.g. Greer in Iowa, USA, and Horton Bluff in Key words: neotype, , species definition.

Rhizodontids (Pisces: Sarcopterygii) are predatory 2000; Parker et al. 2005). However, the absence of a fishes, known from Upper Devonian and Carboniferous robust diagnosis of the type species has led to many finds deposits in many localities world-wide. The UK taxa being assigned to new species. Worse, a variety of material (exclusively Carboniferous) were among the first fossil of uncertain taxonomic status (i.e. any material not associ- fishes to be described. They were worked on by, among ated with the distinctive teeth) has been attributed to the others, Louis Agassiz (Agassiz 1844a), Richard Owen genus, and even to the type species. Finally, two species of (Owen 1840) and Thomas Henry Huxley (Huxley 1862). another rhizodontid genus, also known only from isolated Unfortunately, two genera, Rhizodus Owen, 1840 and teeth (Archichthys Hancock and Athey 1870), were trans- Strepsodus Huxley, in Huxley and Etheridge 1865, have ferred to Strepsodus without explanation by Woodward been caught in systematic and nomenclatural confusions (1891). Although the teeth of Archichthys are distinctive in involving other Devonian and Carboniferous sarcoptery- their own right, they do not resemble those of S. sauroides. gians (Megalichthys, Holoptychius and Rhizodopsis; see The first description of a complete specimen of Strepso- Andrews 1982, 1985 for reviews). The use of the name dus (NMS GY 1980.40.36; Andrews 1985) failed to resolve Rhizodus soon stabilised, although over a century passed the situation. Andrews (1985) noted the nomenclatural before the matter was brought to a close by a plenary rul- problems, and used the specimen as the holotype of a ing of the ICZN (Traquair 1878; Thomson 1966; White new species, ?S. anculonamensis (NB: Andrews assigned and Baird 1967; Melville and China 1970). The type species the species to Strepsodus with some hesitation, but this of Strepsodus, S. sauroides (Binney 1841) was based on an has, in practice, been ignored by subsequent authors). isolated tooth with a distinctive sigmoid shape, and raised Although undoubtedly bearing Strepsodus-like teeth, the striae on the enamel. Until very recently, this was the only specimen is extremely small, and is probably a juvenile certain autapomorphy of the genus. Similar teeth have (cf. Andrews 1985). The bones of the are disarticu- been found in Carboniferous deposits in Europe and lated, imperfectly preserved and most cannot be identified North America (e.g. Dawson 1868; Woodward 1890; God- with any certainty. Thus, although it was extremely frey 1989), and more recently in Australia (Johanson et al. informative on some aspects (it was the first rhizodontid

ª The Palaeontological Association 113 114 PALAEONTOLOGY, VOLUME 49 specimen to show the shape of the body), it was unable In January 1865, John Young presented a paper to the to establish any further Strepsodus autapomorphies. Geological Society of Glasgow using the name ‘Rhizodus This chequered history has led to the unusual situation sauroides’ when referring to H. sauroides, thus implying of a long-established genus having no clear diagnosis, no that it was a rhizodontid, not a holoptychiid (published in holotype, but a plethora of species and attributed material Young 1867; see also Armour 1867; Thompson 1867). Tho- in many collections. New material of Strepsodus is being mas Henry Huxley was also interested in H. sauroides;he described from widespread ages and locations (from the provided an analysis of the genus Holoptychius for Ander- Tournaisian to the Moscovian; from the mid-west of son (1859), including a note on the teeth of a specimen in America to the north of Australia, e.g. Godfrey 1989; the Museum of Practical Geology (MPG) in Jermyn Street, Johanson et al. 2000). This raises biogeographical and London, with ‘fine sharp longitudinal ridges’ (Anderson evolutionary questions, but without a clear definition of 1859, p. 71). Some time before 1865, Huxley appears to the genus, these cannot be addressed. Therefore, this have reclassified specimens of H. sauroides in the museum paper gives a review of the history and material, and pro- to a new genus, Strepsodus. They appear without comment poses a ‘best fix’ for the associated problems. under this genus (but with no species name) in the cata- To aid the comparison of the relative age of sites yield- logue of the museum he published in that year (Huxley and ing Strepsodus remains, I will use international Epoch and Etheridge 1865; cf. Young 1866a, p. 319 ‘…named Strepso- Stage names (Harland et al. 1990), giving the UK equival- dus in the Jermyn Street Collection and Catalogue’). Unfor- ent in parentheses where applicable. tunately, although the MPG catalogue gave accession numbers for each specimen of Strepsodus, they are no lon- ger in the collections of the British Geological Survey (the BRIEF HISTORICAL REVIEW successor of the MPG). At this time, Huxley and Young were also both involved in a committee examining the fos- Binney (1841) described material collected from the sils of the North Staffordshire coalfield. Its report was read Pendleton coalfield [mostly middle Moscovian (Westpha- at the British Association for the Advancement of Science lian B) deposits], which at the time was being mined at meeting in Birmingham in September 1865. The main various pits between Pendleton (Lancashire, England) and report (Egerton et al. 1866) and a supplement by Young Worsley (Greater Manchester, England); OS Landranger (Young 1866a) both used the name Strepsodus with a brief grid references SD757395 and SD747013, respectively. A but unequivocal description of the sigmoid teeth. Young number of specimens, including some distinctive sigmoid (1866b) later elaborated on the separation of Holoptychius teeth with longitudinal ridges on the enamel (Text- sauroides into two genera: Rhizodopsis (not a rhizodontid) fig. 1A; see description, below), were attributed to a new and Strepsodus. He included in Strepsodus some teeth (and species of the porolepiform Holoptychius, H. sauroides associated scales) previously published by Binney (1841), Binney, 1841, p. 165. He then apparently passed the Williamson (1849) and Kirkby and Atthey (1864). Young material on to the leading fish palaeontologist of his day, (1866b) is frequently (and incorrectly) given as the author- Louis Agassiz, so that it could be included in his ‘Recher- ity for the name Strepsodus, even though Young himself ches’. Unfortunately, these fossils are now lost: Jackson’s (ibid., p. 602) clearly cited Huxley. assertion (Jackson 1952) that they are in the collections of Soon after this, Albany Hancock and Thomas Atthey Manchester Museum is incorrect, and my investigations published on large fish remains, including teeth with fine of the literature and other collections have produced no striae, from Moscovian (Westphalian B) deposits in the leads as to its location. Agassiz published the name only coalshales at Newsham, Northumberland, England (Text- (Agassiz 1844a, p. 180; cf. Brown 1890) and gave the type fig. 1E–F, Table 1). At first they assigned it to Rhizodus locality incorrectly as Edinburgh. Meanwhile, Portlock hibberti (Hancock and Atthey 1868) but later established (1843) described and figured some material from Tour- a new genus and species, Archichthys sulcidens Hancock naisian limestones in Londonderry, Northern Ireland, and Atthey, 1870 (see also Hancock and Athey 1871). under an Agassiz manuscript name, Holoptychius portlocki. Grossart (1869, p. 100), working on a scale of Holoptych- However, although the teeth had fine, raised striae on the ius portlocki, noted that it ‘…has now been associated enamel, they were stout and did not resemble those of with a tooth called Strepsodus’. Traquair (1878, 1881) H. sauroides (Text-fig. 1B–C; see description below). In transferred H. portlocki to Archichthys (because of the fact, Portlock (1843, p. 463) noted that they were similar similarity of the teeth), but retaining its specific identity. to those of Rhizodus. Agassiz (1844a, p. 180) eventually Finally, Woodward and Sherborn (1890) listed Archichthys published the name, but did not describe the material. as a possible synonym of Strepsodus, and Woodward However, several subsequent authors followed Portlock in (1891) used the names Strepsodus portlocki and S. sulci- associating the material with Rhizodus (e.g. M‘Coy 1854; dens. Neither publication gave any justification for the Salter 1861; Young and Armstrong 1871). synonymy, but the idea was generally accepted for the JEFFERY: CARBONIFEROUS FISH GENERA 115

A E

BC

D

F

TEXT-FIG. 1. Type material of Strepsodus sauroides, Archichthys portlocki and Archichthys sulcidens. A, isolated tooth, holotype of Strepsodus sauroides, reproduced from a woodcut in Binney (1841, fig. 7). Size unknown because specimen is now lost. B, isolated tooth BGS GSL 28500, part of the type series of Archichthys portlocki, reproduced from a woodcut in Portlock (1843, pl. 13, fig. 8). C, isolated tooth BGS GSL 1301, part of the type series of A. portlocki, reproduced from a woodcut in Portlock (1843, pl. 13, figs 1–3). D, three scales, part of the type series of A. portlocki, reproduced from a woodcut in Portlock (1843, pl. 13, fig. 9). E–F, anterior part of left dentary, NEWHM G59.64, holotype of Archichthys sulcidens. Abbreviations: symphys. tusk, symphysial tusk of dentary; marg. teeth, marginal teeth of dentary; ?overlap id1, probable overlap surface for the first infradentary. next 100 years. Andrews (1985) noted that the teeth of of Natural History, Smithsonian Institution, Washington, USA; these two species were quite distinct from those of the YPM, Yale Peabody Museum, New Haven, USA. type species S. sauroides, and that their inclusion into the genus might be invalid. REVIEW OF COMPARATIVE MATERIAL Abbreviations of institutions. AMF, Australian Museum, Sydney, Australia; AMNH, American Museum of Natural History, New The problem of establishing reliable autapomorphies for York, USA; BGS GSL and GSM, Collections of the British Geo- Strepsodus is exacerbated by other factors. Sites in the UK logical Survey, Keyworth, UK; CPC, Commonwealth Palaeonto- that have yielded rhizodontid remains frequently contain logical Collection, Bureau of Mineral Resources, Canberra, more than one genus or species, leading to difficulty in Australia; GGM, State Geological Museum, Moscow, Russia; assigning isolated bones. The best-documented cases (but GLAHM V, Hunterian Museum Collection, Univer- by no means the only) are Newsham, Northumberland, sity of Glasgow, UK; GRMP, Central Geological Research England [Strepsodus sauroides and Strepsodus (¼Archicht- Museum, Saint Petersburg, Russia; MCZ, Museum of Comparat- ive Zoology, Harvard, USA; MM LL, Manchester Museum Palae- hys) sulcidens] and Gilmerton, Edinburgh, Scotland ontological Collection, UK; NEWHM G, Hancock Museum (Screbinodus ornatus and Rhizodus hibberti). These rhizo- Geological Collections, Newcastle, UK; NHM P, Natural History dontids lived in the freshwater lakes and rivers of a ‘coas- Museum, London, UK; NMS GY, National Museum of Scotland tal alluvial plain’ (see e.g. Boyd 1984; Clarkson 1985; Geological Collections, Edinburgh, UK; NMV P, Museum of Whyte 1994). This was an extremely dynamic palaeoenvi- Victoria, Melbourne, Australia; OSU, Orton Geological Museum, ronment, with constantly changing drainage patterns and Ohio State University, USA; PIN, Palaeontological Institute of sea levels. Thus, the sites (usually mines and quarries) the Russian Academy of Sciences, Moscow, Russia; RM, Redpath often cut through many beds, representing many different Museum, McGill University, Montreal, Canada; SME, Sedgwick palaeoenvironments. Each rhizodontid species must have Museum of Geology, Cambridge, UK; USNM, National Museum 116 PALAEONTOLOGY, VOLUME 49

occupied a separate palaeoenvironmental niche, removed from the others by space or time, but the details of the separation are difficult to ascertain. This problem would be reduced if good collection records were kept [see, for

and example, the records for the Foulden site (Wood and remains reported Strepsodus Rolfe 1985)]. Unfortunately, the majority of rhizodontid material in UK collections was discovered in the nine-

. 2000). teenth century and little documentation exists. In most Strepsodus

et al cases, the sites are no longer accessible (for example, mines and quarries have closed), preventing further sys-

Strepsodus sauroides tematic surveying of the location of rhizodontid remains. These problems mean that in most cases isolated bones can only be assigned to a particular genus with caution.

. A ‘large form’ also reported (Andrews For example, the majority of substantial remains of reported (Johanson teeth reported (Godfrey 1989). reported (e.g. Hancock and Atthey 1870). Strepsodus sauroides come from the Hannah Pit colliery in Newsham, Northumberland (see Table 1), which closed

. 1977). in 1914 (Boyd 1984). This site also produced the majority Strepsodus Strepsodus of the material of Strepsodus [¼Archichthys] sulcidens, and et al (Dawson 1868). so any fossils not closely associated with the distinctive teeth of either species can only be assigned provisionally. To allow a review of the status of Strepsodus and Delta fan. Freshwater lake with surrounding(Andrews swamps. Strepsodus anculonamensis Delta fan. 1985). Brackish, restricted marine embaymentwith with the cyclical sea communication (Tibert and Scott 1999). Type locality of Archichthys sulcidens hardingi Archichthys, the following sections present the material attributed to each species. I restrict the review to material that could be used for comparative purposes within genus Strepsodus and Archichthys. For example, S. anculonamen- sis is by far the best-known species of Strepsodus (known from complete specimens) but many aspects of its anat- omy are of no use in delimiting species, since they are unknown in any other species.

Archichthys the Ducabrook Formation. with some coal shales. Bickett Shales of theChunk Mauch Group. of the Blue Beach Member. Low Main Seam Coalshale. Ox-bow lake within a coal-swamp.

This genus was essentially defined on the basis of its tooth morphology. Almost nothing is known besides this, except an isolated dentary and some more or less dubi- ously assigned scales. mid-Visean. Conglomerate and sandstones of Pendleian. Lochgelly Blackband Ironstone Ivorian (TN3). Berwickshire Cementstones.Brigantian– Arnsbergian. Shallow flood-plain lake, with seasonal flooding. Type locality of Ivorian.Kashirskian (Westphalian B). Shales and ferruginous limestone Strepsodus [¼Archichthys] portlocki Portlock, 1843 (ex Agassiz MS). The type series consists mostly of scales and teeth (now in the collections of the British Geological Survey; BGS GSL 1301–4 and GSM 28500–1) from Car- boniferous limestones in Londonderry, Northern Ireland, which are approximately Ivorian in age (upper Tournai- sian; Dr M. Duncan, pers. comm. 2003). Part of a tetra- pod mandible was also included in the series: BGS GSM 28498 (see Clack and Ahlberg 2004). Very little other material has been attributed to the species. The scales Ducabrook Station, Queensland, Australia. Cowdenbeath, Fife, Scotland. Berwickshire, Scotland. Monongalia County, West Virginia, USA. Horton Bluff, Joggins, Nova Scotia, Canada. Northumberland, England. resemble those of Rhizodus hibberti, but are not well pre- served and bear no diagnostic characters (Text-fig. 1D).

Key sites referred to in this paper. The two tooth crowns (BGS GSL 1301 and GSM 28500) are very similar. BGS GSM 28500 (the best preserved) is 38 mm tall as preserved with the tip snapped off, and Table 1. Ducabrook Middle Paddock, Site nameDora Location Dora opencast site near Age Lithology Palaeontology Foulden Crooked Burn,Greer Foulden, Deckers Creek near Greer, Horton Bluff Newsham Hannah Pit, Newsham, would probably have been 40–45 mm tall when complete JEFFERY: CARBONIFEROUS FISH GENERA 117

(Text-fig. 1B). The root is missing, and it is likely that to its end. This gives the crown a distinctive ‘woven’ texture the tooth had been shed prior to fossilisation, with much (Text-fig. 2A–B), not seen in Rhizodus or Screbinodus. of the dentine resorbed (a process described for Rhizodus; Cruickshank 1968). The base of the crown has 16–18 pli- Strepsodus [¼Archichthys] sulcidens Hancock and Atthey, cations, an indication of the presence of plicidentine 1870. The holotype of this species (NEWHM G59.64; (known to be present in other rhizodontids, and probably Text-fig. 1E–F) is a left dentary from the Kashirskian a primitive characteristic of all stem-, e.g. Schul- (Westphalian B) Low Main Seam Coalshales from the tze 1969, 1970; Long 1989; Jeffery 2003). The tooth is Newsham coal pits (see Table 1). Its taxonomic signifi- gently recurved, similar to the symphysial tusks of Rhizo- cance was forgotten until June 1997, and so it does not dus hibberti and Screbinodus, and much more robust than appear in the Hancock Museum’s catalogue of type those of Strepsodus sauroides. Unlike Rhizodus and Scre- material (Newman et al. 1996). The only material, besides binodus, there are no lingual or labial thickenings (some- a few isolated teeth, comes from Newsham, and almost times inaccurately referred to as ‘cutting edges’ in older all seems to have been collected in the mid-nineteenth literature); as a result the crown is oval, rather than century by Albany Hancock and Thomas Atthey. lenticular, in cross-section. The holotype was accurately described by Hancock and The most distinctive feature is the fine, vertical striae, Atthey (1870). It is preserved in external (labial) view and raised slightly above the surrounding enamel. These sur- is still mostly covered in matrix, although what is visible of round the base of the crown, and also cover its lingual the dermal ornament appears similar to that of Strepsodus (concave), mesial and distal faces, but not the labial (con- sauroides. It resembles the dentary of Rhizodus hibberti in vex) face. They are densely packed, mostly being separated size and shape, and includes a large, plicated symphysial by less than the width of an individual striation. It is diffi- tusk and the bases of nine marginal teeth. The coronoids cult to trace the course of an individual striation, but they and infradentaries are not preserved. The symphysial tusk appear to branch very regularly or at least individual stria- is clearly similar to the teeth of Archichthys portlocki (espe- tions often taper away with a new striation starting adjacent cially BGS GSM 28500). It is oval in cross-section, and

D A B C

TEXT-FIG. 2. Comparison of the teeth of Strepsodus and Archichthys. A–B, Archichthys tooth BGS GSL 1513 in labial view. The irregular striae give a ‘woven’ texture to the enamel. They surround the base of the crown, and rise up the mesial, distal and lingual faces of the crown, but not the labial face. This specimen lacks the tip of the crown, but all known complete examples are simply recurved, not sigmoid. The base of the crown preserves the foldings caused by plicidentine. C–D, Strepsodus NHM P11542 in lateral view. Note the more slender proportions than Archichthys teeth and the sigmoid shape (recurved at the base, reverse-curved at the tip). The regular, well-defined striae surround the very base of the crown and rise up the mesial, distal and lingual faces of the crown, but are absent from the labial (convex) face and the reverse-curved tip. This specimen is broken around the base (very common in isolated Strepsodus teeth) and so does not preserve the external foldings caused by plicidentine. This tooth was probably shed prior to death, as the dentine appears to have been partially resorbed (indicated by the collapsed pulp cavity). Abbreviations: enam. striae, enamel striae; plic. dent., folding caused by plicidentine; pulp cav. collapse, furrow caused by collapse of pulp cavity; recurv., recurved part of tooth; reverse curv., reverse-curved part of tooth. C and D after Jeffery (2003, fig. 11). 118 PALAEONTOLOGY, VOLUME 49 covered in fine, raised striae (although this is not shown in Strepsodus anculonamensis Andrews, 1985. This species, Portlock’s 1843 woodcuts). In some of the basal areas of from the Ivorian (Tn3) Foulden site in Scotland (see the crown the striae are tightly packed and very straight. Table 1), is currently the best known species of Strepsodus, However, tipwards of these areas, the striae become shorter and along with Gooloogongia loomesi from the Upper Devo- and less regular, giving the ‘woven’ texture described for nian of Canowindra, Queensland, Australia (Johanson and Archichthys portlocki. It is possible that the teeth of A. port- Ahlberg 1998, 2001) it is the best known rhizodontid. This locki had basal areas of straighter striae, but that they have is also by far the smallest rhizodontid species; a complete not been preserved. Isolated teeth from Newsham (e.g. individual (the holotype NMS GY 1980.40.36) is only about NEWHM G183.11, G40.52, G59.64 and SME 4741) show 350 mm long, and none of the assigned material comes that the proportion of the crown covered in the straighter from significantly larger individuals. Although small, the striae is variable, but that the more tipward parts of the teeth resemble those of S. sauroides. The species was distin- crown always bear the irregular striae, and have a ‘woven’ guished from S. sauroides by the relatively smaller size of texture. Most of the other isolated teeth attributed to the depressed posterior flange of the cleithrum, and by the Archichthys sulcidens (e.g. SME 4742, MM LL 2397, BGS presumed unconstricted . However, the small GSL 1513 and 1543, and BGS GSM 4551) preserve only the size of the material attributed to this species makes it highly more tipward parts of the crown. probable that it all belongs to juvenile individuals. This Besides the holotype and isolated teeth, the best pre- means that the two autapomorphies could in fact represent served specimen attributed to Archichthys sulcidens is growth-stages (as Andrews 1985 herself noted). Further- NEWHM G186.68b. The specimen consists of a single more, well-preserved specimens attributed to adult Strepso- block with several badly damaged bones, including part dus show a range of variation in the depressed posterior of a right mandibular ramus, bearing two symphysial flange (cf. Jeffery 2001; Parker et al. 2005). In life, it is tusks (a functional tusk and its replacement twin) and thought that part of the pectoral musculature inserted into marginal teeth proportionally smaller than those of Rhizo- the flange, and so a fair amount of variation might be dus hibberti, Screbinodus ornatus and Strepsodus sauroides. expected, depending on the individual’s size, age and A damaged coronoid fang is visible (probably from the strength. second coronoid) and a probable premaxillary tusk. Other than this, none of the bones on the specimen can be Strepsodus arenosus Hay, 1900. This species is based on identified with any certainty. the impression of a scale (USNM 4855) from the ‘Lower Tooth morphology suggests that most of the remaining Carboniferous’ of Blair County, Pennsylvania, USA. The articulated or associated specimens attributed to Archicht- holotype could well be a rhizodontid scale, but scales are hys sulcidens actually belong to large individuals of Streps- notoriously unreliable taxonomic markers. Potential vari- odus sauroides, also common at Newsham (e.g. NEWHM ation of scale morphology with size and age across differ- G7.17, G17.46, G59.66 and G59.65). ent regions of the body and different preservational environments make it nearly impossible to be sure of the specific identity of a rhizodontid scale, especially if found Strepsodus in isolation (contra Andrews 1985). There is little or no other material attributed to S. arenosus. Strepsodus sauroides (Binney, 1841, pars). The morphol- ogy of the teeth of Strepsodus sauroides was described in Strepsodus brockbanki Davis, 1891. This species was esta- detail by Jeffery (2003). Like those of Archichthys, they are blished on the basis of material discovered in Moscovian oval in cross-section and bear striae. However, they (Westphalian B) limestone, at Levenshulme near Man- are easily distinguished from Archichthys teeth, as they are chester, England, apparently with more robust teeth than relatively taller and more slender; the striae are longer, S. sauroides. The single specimen of this species (the more regular, much more sharply defined and less densely holotype MM LL222) is too fragmentary to identify any packed; the striae are restricted to the lingual (concave) rhizodontid characteristics. face and parts of the mesial and distal faces; the striae are much straighter than recurved crown, causing striae on Strepsodus dawsoni Hay, 1900. This species was based on a the lingual face at the base of the crown to run onto the single scale (RM 3076) from the coal shales of Pictou, Nova mesial or distal faces toward the tip of the crown; the Scotia, Canada, dated from Podolskian to early striae never degenerate to give the ‘woven’ texture (see Myachkovskian (c. Westphalian C to lower Westphalian D; Text-fig. 2C–D). Many teeth are recurved, but some Dr G. Yeo, pers. comm. 2003). It was originally figured by (possibly only the symphysial and premaxillary tusks) are Dawson (1868, fig. 56b), along with a tooth, as sigmoid, recurved along most of their length, with a slight Rhizodus lancifer Newberry, 1856 (now Anthracosaurus lan- reverse-curve at the very tip. cifer; see Romer 1963). Hay (1900) stated that it was similar JEFFERY: CARBONIFEROUS FISH GENERA 119 to the scales attributed to S. hardingi by Dawson (1868), (Table 1; Andrews et al. 1977; Andrews 1985, p. 80). A but decided that its appearance in the much younger Pictou dentary (NMS GY 1975.48.31) and two postparietals coalfields warranted the establishment of a new species. (NMS GY 1978.4.16 and 17; Text-fig. 3B–C, E–F) are very The scale certainly appears to be rhizodontid, but cannot similar to material attributed to S. sauroides from New- be assigned to a genus with any certainty (see above). How- sham (Text-fig. 3A, D; Jeffery 1999, 2001). ever, its occurrence in the Pictou coal shales is notable as the youngest known rhizodontid fossil; the next youngest Foulden large form. This material from the Ivorian fossils are Kashirskian (Westphalian B, e.g. Newsham). (TN3) Foulden site in Scotland (Table 1) was described There is little or no other material attributed to S. dawsoni. by Andrews (1985), but not named by her. It includes cleithra, clavicles and jaw material. Three isolated teeth Strepsodus hardingi (Dawson, 1868). This was originally (NHM P13124, 13127 and 13128) are similar to Archicht- described as a species of Rhizodus, but Woodward (1890, hys sulcidens (White 1927; Jeffery 2003). A partial dentary 1891) transferred it to Strepsodus. The type series (from the (NMS GY 1980.40.46) is similar to material attributed to Ivorian of Horton Bluff, Nova Scotia, Canada; Table 1) is S. sauroides (Jeffery 2003). One clavicle (NMS GY close in age and palaeogeographical location to the Foulden 1980.40.49) has a quadrangular extension to the base of site of Scotland (Table 1; cf. S. anculonamensis and the the clavicular spine not seen in any other rhizodontid ‘Large form’). It includes part of a mandible which, along (Andrews 1985, fig. 11c). However, given the variation of with newly discovered material, is currently under study by the pectoral girdle (Parker et al. 2005) the significance of Martin Brazeau at the Redpath Museum. It appears to pos- this is not clear. sess several autapomorphic characters that may distinguish it from Strepsodus (M. Brazeau, pers. comm. 2004). Greer Rhizodontid. This material is from Greer, West Virginia, USA (Table 1; Godfrey 1989). Knowing the Strepsodus minor Traquair, 1890. Almost no material has taxonomic problems of rhizodontids, Godfrey (1989) been attributed to the species beyond the type series from refrained from assigning the material to a species. How- the Asbian of Pitcorthy (or Pitcornie), Dunfermline, Scot- ever, an isolated tooth (MCZ 13889; Godfrey 1989, pl. 1) land (Traquair 1890; Jeffery 2003). A tooth belonging to clearly belongs to Strepsodus (cf. Parker et al. 2005). the type series (NMS GY 1881.43.42) is indistinguishable Rhizodontid remains from a slightly younger site in Ken- from the most densely striated of the teeth of S. sauroides. tucky, USA, have not been described in detail (see Ches- nut et al. 2002). However, the Greer and Delta sites have Strepsodus striatulus Traquair, 1882. This species was similar vertebrate faunas (see Schultze and Bolt 1996), so established on the basis of isolated teeth collected from Strepsodus might also be present at Delta. the lower Moscovian (Westphalian B) of Burghlee (¼Bor- ough Lee), Loanhead, Scotland. The limited amount of Devonian material. Rhizodontid remains from two material attributed to it was reviewed by Jeffery (2003). Upper Devonian localities have been attributed to Strepso- None of it ‘is distinguishable from S. sauroides, if some dus: the Ko¨pru¨lu¨ shales in Turkey (Janvier et al. 1984) degree of variation in tooth recurvature is accepted’ and the in Colombia (Janvier and Vil- (Jeffery 2003, p. 266). larroel 2000). Although both sets of remains are rhizo- dontid, neither belongs to Strepsodus (see Jeffery 2001, 2003) and they will not be further considered here. Material not currently assigned to a species

Ducabrook Strepsodus. Material from the mid-Visean Strepsodus and Onychodus Ducabrook site in Queensland, Australia (Table 1), has been described by Johanson et al. (2000) and Parker et al. Teeth of Strepsodus have occasionally been mistaken for (in press). Teeth similar to those of S. sauroides have been the Carboniferous occurrences of Onychodus, an onycho- discovered, and the pectoral girdle and fin are dontid sarcopterygian that also bears raised striae on sig- comparable (given the variation seen in the UK material). moid teeth (see Jeffery 2003, p. 266 for a discussion). However, although confident that the material could be However, the two are easy to tell apart; onychodontid assigned to Strepsodus, Parker et al. (2005) were aware of teeth lack sizeable roots and have only partial dentine the taxonomic difficulties besetting the genus and did not folding (e.g. Schultze 1969, p. 126, fig. 26). Also, assign the material to any species. the striae are usually more densely packed and follow the recurvature of the crown more closely than those of Cowdenbeath material. This material comes from the Strepsodus. Besides these mistakenly identified teeth, ony- Pendleian Dora opencast site near Cowdenbeath, Scotland chodontid remains are restricted to the Devonian. 120 PALAEONTOLOGY, VOLUME 49

A BC D EF

TEXT-FIG. 3. Postparietals of Strepsodus from the UK. A, D, left postparietal, NEWHM G18.93 from Newsham, Northumberland [Kashirskian (Westphalian B)]. B, E, left postparietal, NMS GY 1978.4.16 from the Dora opencast site (Pendleian); C, F, right postparietal NMS GY 1978.4.17, also from Dora. Despite the age difference (over 20 myr), these postparietals all show a possible autapomorphy of Strepsodus, the deep, angled overlap surface for the lateral extrascapular. All specimens are in external view. Abbreviations: overlap lat. extrascap., overlap surface for lateral extrascapular.

TAXONOMIC CONCLUSIONS exist that can separate material of the two species (Archichthys portlocki and A. sulcidens). I therefore suggest Defining species in the fossil record is often difficult, that, unless and until such autapomorphies are deter- especially when only a limited number of fragmentary mined, the material be treated as a single species. The specimens are available, as is the case with Strepsodus and name Archichthys portlocki has priority. Archichthys. The most useful definitions in such circum- It should also be noted here that, although many bones stances are character-based (e.g. phylogenetic species con- attributed to Archichthys certainly resemble those of rhizo- cepts; Cracraft 1983; Nixon and Wheeler 1990; Baum and dontids (e.g. the dentary NEWHM G59.64), no specimen Donoghue 1995), demanding that a species possess at has yet been found that shows any unequivocal rhizodontid least one unique, derived character (an autapomorphy). synapomorphies (e.g. elongated basal lepidotrichial seg- Under all taxonomic systems, the criteria for placing a ments, ‘reverse overlap’ of the clavicle and cleithrum, a species in a separate genus are arbitrary, but are usually a depressed posterior flange of the cleithrum, narrow articu- pragmatic indication of a more ‘distant’ relationship. lation between median extrascapular and postparietals; cf. In the case of Strepsodus, the delimitation of species Andrews and Westoll 1970; Andrews 1973, 1985; Cloutier can be explored in the form of a thought experiment: if and Ahlberg 1996; Johanson and Ahlberg 1998, 2001; Jeff- all the specimens of Strepsodus, except type material, had ery 1999, 2001). A few specimens (in particular the dentary their museum labels removed, which specimens could be NEWHM G59.64) indicate membership of the stem tetra- successfully re-classified with reference to the type mater- pod lineage (of which rhizodontids form a basal plesion), ial? The review of material presented above allows several but cannot rule out the possibility that Archichthys might taxonomic conclusions, listed below. belong to another stem tetrapod group.

The status of Archichthys Species of Strepsodus

It is clear that the two species formally assigned to Type specimens for S. sauroides. As noted above, the holo- Archichthys are easily distinguished from the type species type of S. sauroides (Binney 1841, fig. 7; reproduced in of Strepsodus (S. sauroides) based on tooth morphology. Text-fig. 1A) is lost, as are the specimens that Huxley first Woodward and Sherborn (1890) and Woodward (1891) named Strepsodus (Huxley and Etheridge 1865, p. 145). A never gave reasons for their inclusion in Strepsodus.I type specimen is needed in order to clarify the status of therefore agree with Andrews (1985) in asserting that other Strepsodus species. Three teeth from Binney’s collec- Archichthys is a valid genus. However, no autapomorphies tion are stored in the Sedgwick Museum, Cambridge JEFFERY: CARBONIFEROUS FISH GENERA 121

(SME 4721, 4723 and 4746). I propose that one of these other characters that may be autapomorphies of specimens, SME 4721 (Text-fig. 4), be nominated the S. sauroides (sensu stricto), although this cannot be con- neotype under ICZN Article 75 (Ride et al. 2000). This firmed until more is known of the morphology of other specimen was collected from Moscovian (Westphalian) rhizodontids. The first concerns the overlap of the lateral coal shales at Longton, Staffordshire, England (part of the extrascapulars with the postparietals. Postparietals from Pottery Coalfield of North Staffordshire; OS Landranger Dora, Scotland (NMS GY 1978.4.16 and 17), found in grid reference SJ 905432) approximately 70 km south of association with Strepsodus sauroides jaw material bear a where the holotype was collected (at an unknown loca- distinct ‘notch’ in the posterolateral margin, and a large tion within the Pendleton coalfield; see above). However, overlap surface where the lateral extrascapular articulated it was clearly identified as ‘Holoptychius sauroides’ by Bin- (Text-fig. 3B–C, E–F). This matches isolated postparietals ney himself, as the specimen still bears his label. There- from Newsham, England (e.g. NEWHM G18.93; Text- fore, this specimen best fulfils the requirements of ICZN fig. 3A, D), attributed to Strepsodus sauroides and con- Article 75.3.5 that the neotype be ‘consistent with what is trasts with the postparietals of Screbinodus (e.g. NMS known of the former name-bearing type’. GY 1963.16.18) where the overlap is straight with a shal- low overlap (Andrews 1985, fig. 7c). The degree of over- Nomina dubia. On the basis of inadequate type material, lap is unknown in Gooloogongia and Barameda. the following species should be considered nomina dubia: However, AMF100073A shows that the articulation was Strepsodus arenosus Hay, 1900; S. brockbanki Davis, 1891; fairly straight in Gooloogongia (Johanson and Ahlberg S. dawsoni Hay, 1900. 2001) and NMV P160880 shows that the articulation was gently curved in Barameda (Long 1989, although Junior synonyms. On the basis of the type material, the larger rhizodontid material from the same location following species should be considered junior synonyms suggests that this specimen is a juvenile, and thus the of Strepsodus sauroides: S. anculonamensis Andrews, 1985; articulation may have altered with growth). This S. minor Traquair, 1890; S. striatulus Traquair, 1882. autapomorphy must be regarded as provisional, in par- S. anculonamensis includes the most complete specimen ticular because there is a possibility that the postparietals of Strepsodus (NMS GY 1980.40.36). It shows teeth that from Newsham in fact belong to Archichthys portlocki are clearly similar to those of S. sauroides but, as dis- (sensu stricto). Unfortunately, the postparietals of S. an- cussed above, it possesses no autapomorphies which culonamensis (sensu lato) are preserved in internal view, could not equally be explained by the probable juvenile and the extrascapulars are not preserved at all, prevent- state of all the specimens, bearing in mind that specimens ing an assessment of the oldest Strepsodus material. Also, were found in the same beds as the ‘Foulden large form’, nothing is known of the extrascapulars of Rhizodus; a Strepsodus with a body-length of perhaps 3Æ5m Andrews’ interpretation of an isolated Rhizodus postpari- (Andrews 1985, p. 92). I have not had the opportunity to etal (Andrews 1985, fig. 7d) must be treated with cau- examine the material of S. hardingi personally; an assess- tion. ment of the status of this species is being undertaken by Other potential autapomorphies have been suggested Martin Brazeau at the Redpath Museum. by Johanson et al. (2000) and Parker et al. (2005). The latter review the morphology of isolated humeri Other material. On the basis of published material, the found at Ducabrook (Australia) and Greer (US), both following are suggested to belong to Strepsodus sauroides: sites that have yielded only Strepsodus-like teeth. They the ‘Cowdenbeath material’ from the Dora opencast site, identified six characters that are shared with isolated Scotland (Table 1; cf. Andrews et al. 1977; Andrews humeri attributed to Strepsodus from Newsham (Eng- 1985); the ‘Foulden large form’ from Foulden, Scotland land), and which are not seen in the other well-known (Table 1; Andrews 1985), with the exception of three iso- rhizodontid humeri [Aztecia (Johanson and Ahlberg lated teeth (NHM P13124, 13127 and 13128) hereby 2001), Sauripterus (Davis et al. 2004), Rhizodus (Jeffery assigned to Archichthys portlocki (sensu stricto). I have not 2001) and Barameda (Garvey et al. 2005)], and may had the opportunity to examine the specimens from therefore be Strepsodus autapomorphies. These characters Greer, USA, or Ducabrook, Australia, personally (Table 1; are: (1) ulnar and radial condyles are offset from one Godfrey 1989; Johanson et al. 2000; Parker et al. 2005). another (i.e. they face distally in slightly different direc- However, unless and until autapomorphies are discerned tions); (2) humeral ridge is shallow in comparison with that can reliably distinguish this material from S. sauro- entepicondyle and pectoral process; it forms a deep notch ides, I suggest they are treated as S. sauroides. between the two; (3) pectoral process is large (a third to a half of the area of the entepicondyle); (4) pectoral pro- Possible autapomorphies of Strepsodus sauroides. Besides cess is separated from the dorsal ridge by the pectoral the tooth morphology described above, there are some ridge; (5) dorsal ridge runs toward the ulnar condyle; (6) 122 PALAEONTOLOGY, VOLUME 49 dorsal ridge forms a V-shaped structure with the ectepi- SYSTEMATIC PALAEONTOLOGY condyle; and (7) entepicondyle is long and gracile (com- pared with Rhizodus). The following diagnoses use some details from Jeffery (1999, Johanson et al. (2000, p. 165) suggested that the 2001, 2003), with additional information from Johanson et al. Ducabrook cleithra and cleithra of S. sauroides from (2000) and Parker et al. (2005). Newsham, England, share a ‘well developed pectoral Owing to the complex taxonomic history of Strepsodus, incision and [depressed] posterior flange’, the depressed Archichthys and other Rhizodontida (sensu Andrews and Westoll posterior flange having a rounded shape (Johanson 1970), older literature and museum catalogues may use a bewil- dering array of different names for the same . To help et al. 2000, fig. 2). This may be correct, although more overcome this problem, I append a check-list of published must be known of the individual variation caused by names, including for each type material, a brief history and the size, age, muscle distribution and usage before this can current taxonomic status. be applied with full confidence (Ducabrook material is from relatively small individuals, when compared with much of the material from the UK). For example, one SARCOPTERYGII Romer, 1955 particular cleithrum from Newsham attributed to Streps- RHIZODONTIDA Andrews and Westoll, 1970, p. 463 odus sauroides (NEWHM G19.41) bears a very robust depressed posterior flange, more similar to that of Rhiz- Genus STREPSODUS Huxley, in Huxley and Etheridge 1865, odus than to those of several of the other cleithra, also p. 145 from Newsham, attributed to S. sauroides. This speci- men may demonstrate individual or sexual variation or Type species. Strepsodus sauroides (Binney, 1841). a growth sequence, or it may even belong to Archicht- hys. Either way, until more data are available, cleithral Diagnosis. Teeth tall and oval in cross-section, with regu- characters must be considered indicative, but not lar, even striae on the lingual, mesial and distal surface of diagnostic. crown. Labial face of crown smooth and free of striae. Striae are much less recurved than the crown as a whole, and thus a striation with a lingual position on the base of Age the crown may take up a mesial or distal position towards the tip of the crown. Each striation is 40–50 lm wide, The time range of a single, consolidated S. sauroides is long regardless of the size of the tooth. There are 6–12 ) (Ivorian–Kashirskian, over 40 myr). It is, of course, highly striae.mm 1. Base of the crown bears placations indicative probable that breeding populations varied in size and dis- of plicidentine (although this characteristic is often tribution over this time (i.e. forming different ‘biospecies’). obscured on teeth preserved in situ, and lost from isolated However, nothing of this is yet clearly discernible in the teeth). Teeth may be recurved or slightly sigmoid. If sig- fossil record. Material from Foulden, Scotland (Ivorian; moid, the reverse curvature is restricted to the apical 10 per Table 1), is morphologically indistinguishable from mater- cent of the crown. On the humerus, the ulnar and radial ial from Dora, Scotland (Pendleian; Table 1), and New- condyles are offset from one another; the humeral ridge is sham, England (Kashirskian; Table 1). shallow in comparison with entepicondyle and pectoral process, forming a deep notch between the two; the pec- toral process large (one-third to half of the area of the ent- Palaeogeography epicondyle); the pectoral process separated from the dorsal ridge by the pectoral ridge; the dorsal ridge runs toward the The Euramerican sites were relatively close (at a similar ulnar condyle and forms a V-shaped structure with the palaeolatitude and spanning less than 60 degrees of pal- ectepicondyle; the entepicondyle is long and gracile. aeolongitude), and most probably formed part of a more Two additional possible diagnostic characters (cleithrum or less contiguous ‘coastal alluvial plain’ (cf. Anderton with small, rounded, depressed posterior flange; postpari- 1985). However, the Australian Ducabrook material is etal with posterolateral notch and large dished overlap area much more distant, and indicates that Strepsodus was for lateral extrascapular) cannot yet be confirmed, but widely spread (presumably across the south-east of Laur- should be considered indicative rather than diagnostic. asia to the north-east of ). This distribution is mirrored in some other groups, notably the whatcheerid tetrapods (cf. Parker et al. 2005). Unfortunately, little or Strepsodus sauroides (Binney, 1841) no data are available for any intervening Carboniferous freshwater or coastal sites (e.g. from the Middle East or 1841 Holoptychius sauroides Binney, p. 165, pl. 5, central Asia). figs 7–7a. JEFFERY: CARBONIFEROUS FISH GENERA 123

1866 Dendroptychius sp. Huxley, in Huxley and Ethe- ridge 1865; Young, p. 601, fig. 7 [cf. Woodward 1891, p. 349]. 1867 Rhizodus sauroides (Binney, 1841); Young, p. 39 [text only]. non 1868 Rhizodus hardingi Dawson, p. 255, fig. 77a–d. 1882 Strepsodus striatulus Traquair, p. 544 [text only]. 1890 Strepsodus minor Traquair, p. 393 [text only]. non 1891 Strepsodus hardingi (Dawson, 1868); Woodward, p. 353 [text only]. 1943 Onychodus ostrovensis von Huene, 1943, p. 100, fig. 1. 1973 Onychodus cf. ostrovensis von Huene, 1943; Schultze, pl. 33, fig. 4 [see Schultze 1993; Jeffery 2003]. non 1984 Strepsodus sp.; Janvier et al., p. 158, pl. 2, figs 9– 12, fig. 9A–F. 1985 Strepsodus anculonamensis Andrews, p. 71, figs 1–3, 5a–e, 6, 7a, 8, 9a–c, 10a, d, 11a, 14c. TEXT-FIG. 4. Isolated tusk SME 4721 from Longton, 1985 Large form; Andrews, p. 83, figs 5g–h, 7b, 9d–e, Staffordshire, England, the proposed neotype of Strepsodus 10e, 11b–e, 12b. sauroides (Binney, 1841). This tooth is sigmoid, shows the non 1985 Large form; Andrews, fig. 12a [¼Archichthys distinctive enamel striae and is apparently similar to the lost portlocki]. holotype: importantly, it is one of the only specimens known 1985 Cowdenbeath material; Andrews, p. 80 [text only]. that was attributed to Holoptychius (¼Strepsodus) sauroides by 1989 Greer rhizodontid; Godfrey, p. 89, figs 1–5. Binney himself (cf. ICZN Article 75.3.5). non 2000 Strepsodus?; Janvier and Villarroel, pp. 750–752, fig. 15. Archichthys portlocki Portlock, 1843 ex Agassiz MS 2000 Ducabrook Strepsodus; Johanson et al., p. 161, fig. 3. 1843 Holoptychius portlocki Portlock, ex Agassiz MS, 2005 Ducabrook Strepsodus; Parker et al., figs 2–3, p. 463, pl. 13, figs 1–13. 4a–c, 5a, 6–8, 10, 11, 14. 1844 Holoptychius portlockii Portlock, 1843 ex Agassiz MS; Agassiz 1844a, p. 180 [text only]. Note that these are only synonyms with taxonomic significance; 1861 Rhizodus portlockii (Portlock, 1843 ex Agassiz MS); see Woodward (1891) for a comprehensive list of early syno- Salter, p. 223 [text only]. 1868 Rhizodus hibberti Owen, 1840; Hancock and Atthey, nyms of S. sauroides (sensu lato). p. 346 [text only; see Hancock and Atthey 1870]. 1870 Archichthys sulcidens Hancock and Atthey, p. 266 Types. Holotype is an isolated tooth from the Pendleton coal- [text only]. field near Manchester, England (now lost); neotype is an isolated 1891 Strepsodus sulcidens (Hancock and Atthey, 1870); tooth, SME 4721, from Longton, Staffordshire, England (Text- Woodward, p. 352 [text only]. fig. 4). 1891 Strepsodus portlocki (Portlock, 1843 ex Agassiz MS); Woodward, p. 353 [text only]. Diagnosis. As for genus (only species). 1903 Strepsodus sulcatus Traquair, 1903, p. 700 [lapsus calami, text only]. Age range. Ivorian (Foulden; Andrews, 1985) to Kashirskian 1927 Strepsodus cf. sulcidens (Hancock and Atthey, 1870); (e.g. Newsham; Boyd 1984), but possibly to the Myachkovskian White, p. 259 [text only]. (Pictou coals; Dr G. Yeo, pers. comm. 2003). 1985 Large form; Andrews, fig. 12a.

Distribution. UK, Northern Europe and North America, Type series. Teeth (BGS GSL 1301–1302 and BGS GSM 28500) Australia. and scales (BGS GSL 1304 and BGS GSM 28501). BGS GSM 28498 and 28502 are excluded here from Portlock’s original type series as they belong to a tetrapod (see Clack and Ahlberg 2004). Material Genus ARCHICHTHYS Hancock and Atthey, 1870 from various localities in Co. Londonderry, Northern Ireland.

Type species. A. portlocki Portlock, 1843 ex Agassiz MS. Diagnosis. As for genus (only species).

Diagnosis. Teeth robust and recurved with fine, irregular Age range. ?Ivorian (?upper Tournaisian) of Londonderry, Nor- striae, giving a ‘woven’ texture. thern Ireland (Duncan 1999) to Kashirskian (Westphalian B) of 124 PALAEONTOLOGY, VOLUME 49

Newsham, Northumberland, England (Hancock and Atthey Museum of Zoology, funded by BBSRC grant 9430 6007. I am 1870). grateful to my supervisor, Jenny Clack, for her help and advice. Many people have assisted me with dating and strati- Distribution. UK. graphy, as well as locating specimens and literature relating to rhizodontids: many thanks to Per Ahlberg, Sarah Finney, Jeff Liston, Neil Clark, Mags Duncan, Bobbie Paton, Steve Tunni- DISCUSSION cliffe and Gary Yeo. I am also indebted to Marcelo Ruta, Jil- lian Garvey, Anne Warren, Zerina Johanson and especially Kate Parker, Martin Brazeau and the late Mahala Andrews for Despite the wide distribution of Strepsodus sauroides in helpful and stimulating discussion. The original manuscript space and time, it should be stressed that splitting the benefited greatly from the comments made by Zernia Johan- taxon into several species serves no useful taxonomic pur- son and Gae¨l Clement. pose, nor does it truly reflect our current understanding of the genus Strepsodus. With so few specimens, mostly comprising isolated scales, teeth and bones, no account REFERENCES can be taken of individual or sexual variation of the ele- ments in question, nor how the variations are distributed AGASSIZ, J. L. R. 1844a. Recherches sur les poissons fossiles in time and space. Also, in purely practical terms, it (Tome 2, Part 2). Imprimerie de Petitpierre, Neuchaˆtel (aux serves little phylogenetic purpose. The Rhizodontida as a frais de l’auteur), 336 pp. whole was not a large radiation; it currently contains only —— 1844b. Monographie des poissons fossiles du Vieux Gre`s seven reliably diagnosed genera (Aztecia Johanson and Rouge ou Syste`me De´vonien des Iles Brittanniques et de Russie Ahlberg, 1998; Barameda Long, 1989; Gooloogongia Johan- (Livraison 2). Jent et Gassmann, Neuchaˆtel et Soleure, 72 son and Ahlberg, 1998; Rhizodus Owen, 1840; Sauripterus pp. Hall, 1843; Screbinodus Andrews, 1985; Strepsodus Huxley, ANDERSON, J. 1859. Dura Den; a monograph of the Yellow Sandstone and its remarkable fossil remains. Thomas Constable in Huxley and Etheridge 1865) with the possibility of a & Co., Edinburgh, 96 pp. few more, for example Archichthys Hancock and Atthey, ANDERTON, R. 1985. Sedimentology of the Dinantian of 1870, and some of the Devonian rhizodontid material Foulden, Berwickshire, Scotland. Transactions of the Royal previously attributed to Strepsodus (Janvier et al. 1984; Society of Edinburgh: Earth Sciences, 76, 7–12. Janvier and Villarroel 2000; cf. Jeffery 2001, 2003). All are ANDREWS, S. M. 1973. Interrelationships of crossoptery- monotypic (see Appendix), and most can be easily distin- gians. 138–177. In GREENWOOD, P. H., MILES, R. S. guished from the other genera. Thus, any separation of and PATTERSON, C. (eds). Interrelationships of fishes. Strepsodus based on currently known material would sim- Zoological Journal of the Linnean Society, 53, Supplement ply create a (more or less reliable) sister species of No. 1, 536 pp. S. sauroides and add little to our understanding of rhizo- —— 1982. The discovery of fossil fishes in Scotland up to 1845 dontid phylogeny. In fact, the limited amount of material with checklists of Agassiz’s figured specimens. Royal Scottish Museum Press, Edinburgh, 87 pp. available for Strepsodus has meant that all recent phylo- —— 1985. Rhizodont crossopterygian fish from the Dinatian of genetic analyses of the Rhizodontida have (quite reason- Foulden, Berwickshire, Scotland, with a re-evaluation of this ably) ignored all species divisions, and treated Strepsodus as group. Transactions of the Royal Society of Edinburgh: Earth a single operational taxonomic unit (e.g. Young et al. 1992; Sciences, 76, 67–95. Johanson and Ahlberg 1998, 2001; Jeffery 1999, 2001, –––– BROWNE, M. A. E., PANCHEN, A. L. and WOOD, S. 2003). However, if reliable specific differences can ever be P. 1977. Discovery of amphibians in the Namurian (Upper identified, it would make more sense to establish new spe- Carboniferous) of Fife. Nature, 265, 529–532. cies rather than attempt to rationalise the old, less reliably –––– and WESTOLL, T. S. 1970. The postcranial skeleton of rhi- defined species with their attendant ‘taxonomic baggage’. pidistian fishes excluding Eusthenopteron. Transactions of the Perhaps the most significant question to ask is what Royal Society of Edinburgh, 68, 391–489. factors allowed a species (or tribe of sister-species) to ARMOUR, A. 1867. ‘Mr. Andrew Armour exhibited–’. Trans- actions of the Geological Society of Glasgow, 2, 69. retain a set of characters unchanged for over 40 million BAIRD, D. 1978. Studies on Carboniferous freshwater fishes. years? This suggests a surprising stability of the Strepsodus American Museum Novitates, 1641, 1–22. niche within the very dynamic physical environment of BALL, H. W., DINELEY, D. L. and WHITE, E. I. 1961. The the alluvial plain. Correlations should be sought with Old Red Sandstone of Brown Clee Hill and the adjacent area. other species, to investigate if potential prey or competi- Bulletin of the British Museum (Natural History) (Geology), 5, tors remained similarly stable over such a long time. 175–310. BARDOCK, D. 1979. Fishes of the Mazon Creek fauna. Acknowledgements. The research for this paper was carried 501–528. In NITECKI, M. H. (ed.). Mazon Creek fossils. out as part of my PhD project at the University of Cambridge Academic Press, New York, 581 pp. JEFFERY: CARBONIFEROUS FISH GENERA 125

BARKAS, T. P. 1873. Illustrated guide to the fish, amphibian, and Prince Edward Island. Second edition. MacMillan, Lon- reptilian, and supposed mammalian remains of the Northumber- don, 694 pp. land Carboniferous strata. W. M. Hutchings, London, 117 pp. DUNCAN, M. 1999. A study of some Irish Lower Carbonifer- BAUM, D. A. and DONOGHUE, M. J. 1995. Choosing ous fish microvertebrates. Unpublished PhD thesis, University among alternative ‘phylogenetic’ species concepts. Systematic of Dublin, 225 pp. Botany, 20, 560–573. EGERTON, P. M. G., HUXLEY, T. H. and MOLYNEUX, BINNEY, E. W. 1841. On the fossil fishes of the Pendleton coal W. 1866. Report of the committee on the distribution of the field. Transactions of the Manchester Geological Society, 1, 153– vertebrate remains from the North Staffordshire coal field. 178. Reports of the British Association for the Advancement of Sci- BOYD, M. J. 1984. The Upper Carboniferous tetrapod assem- ence, 1865, 42–51. blage from Newsham, Northumberland. Palaeontology, 27, ETHERIDGE, R. 1888. Fossils of the British Islands stratigraph- 367–392. ically and zoologically arranged. Volume 1; Palaeozoic, compri- BROWN, W. H. 1890. Dates of publication of ‘Recherches sur sing the , , Devonian, Carboniferous, and les Poissons Fossiles’, par L. Agassiz with a note on Owen’s species. Clarendon Press, Oxford, 468 pp. ‘Odontography’. xxv–xxix. In WOODWARD, A. S. and GARVEY, J. M., JOHANSON, Z. and WARREN, A. SHERBORN, C. D. (eds). Catalogue of British fossil Verteb- 2005. Redescription of the pectoral fin and vertebral column rata. Dulau, London, 396 pp. of the rhizodontid fish Barameda decipiens from the Lower CHABAKOV [KHABAKOV], A. W. 1927. On the remains Carboniferous of Australia. Journal of Vertebrate of the crossopterygians of the Carboniferous of Russia. Paleontology, 25, 8–18. Izvestiya Geologicheskogo Komiteta, 46, 299–309. GODFREY, S. J. 1989. A rhizodontid crossopterygian from CHESNUT, D. R., GREB, S., STORRS, G. W., GARCIA, the Upper Mississippian at Greer, West Virginia. Acta W. J. and BELLAN, J. 2002. Sedimentology and palaeoecol- Musei Reginaehradecensis, Series A: Scientiae Naturales, 22, 89– ogy of a Namurian A tetrapod site. Buffalo Wallo Formation, 98. Western Kentucky. IPC2002, Geological Society of Australia, GROSSART, W. 1869. On the Upper Coal Measures of Lan- Abstracts, 68, 195. arkshire. Transactions of the Geological Society of Glasgow, 3, CLACK, J. A. and AHLBERG, P. E. 2004. A new stem 96–113. tetrapod from the Early Carboniferous of Northern Ireland. HALL, J. 1840. Fourth Annual Report of the Survey of the 309–320. In ARRATIA, G., WILSON, M. V. H. and Fourth Geological District. Assembly Document, Albany Insti- CLOUTIER, R. (eds). Recent advances in the origin and early tute, 50, 389–480. radiation of . Verlag Friedrich Pfiel, Munich. —— 1843. Natural history of New York; geology of New York, CLARKSON, E. N. K. 1985. Palaeoecology of the Dinantian of part IV, comprising the survey of the Fourth Geological District. Foulden, Berwickshire, Scotland. Transactions of the Royal Carroll & Cook, Albany, USA, 683 pp. Society of Edinburgh: Earth Sciences, 76, 97–100. HANCOCK, A. and ATTHEY, T. 1868. Notes on the remains CLOUTIER, R. and AHLBERG, P. E. 1996. Morphology, of some reptiles and fishes from the shales of the Northum- characters, and the interrelationships of basal sarcopterygians. berland Coal-field. Annals and Magazine of Natural History, 1, 445–479. In STIASSNY, M. L. J., PARENTI, L. R. and 266–278. JOHNSON, G. D. (eds). Interrelationships of fishes. Academic —— —— 1870. Note on an undescribed fossil fish from the Press, London, 496 pp. Newsham coal-shale near Newcastle-upon-Tyne. Annals and COPE, E. D. 1897. On new Vertebrata from Illinois, Magazine of Natural History, 5, 266–268. Ohio and Pennsylvania. Proceedings of the American Philosoph- —— —— 1871. Description of a considerable portion of a ical Society, 36, 71–91. mandibular ramus of Anthracosaurus Russelli; with notes on CRACRAFT, J. 1983. Species concepts and speciation analysis. Loxoma and Archichthys. Transactions of the Tyneside Natural- 159–187. In JOHNSTON, R. F. (ed.). Current ornithology. ists’ Field Club, 4, 385–397. Volume 1. Plenum Press, New York, 425 pp. HARLAND, W. B., ARMSTRONG, R. L., COX, A. V., CRUICKSHANK, A. R. I. 1968. Tooth structure in Rhizodus CRAIG, L. E., SMITH, A. G. and SMITH, D. G. 1990. A hibberti Ag., a rhipidistian fish. Palaeontologica Africana, 11, geologic time scale 1989. Cambridge University Press, Cam- 3–13. bridge, 263 pp. DAVIS, J. W. 1891. On the discovery of a new species of fossil HAY, O. P. 1900. Descriptions of some vertebrates of the fish (Strepsodus brockbanki) in the Upper Coal Measures Lime- Carboniferous Age. Proceedings of the American Philosophical stone of Levenshulme, 6 Group, from the railway cutting at Society, 39, 96–115. Levenshulme, near Manchester. Memoirs and Proceedings of HIBBERT, S. 1835. Memoir on the fresh-water limestone of the Manchester Literary and Philosophical Society, 4, 427–429. Burdiehouse, in the neighbourhood of Edinburgh, belonging DAVIS, M. C., SHUBIN, N. and DAESCHLER, E. B. 2004. to the Carboniferous group of rocks; with supplementary A new specimen of Sauripterus taylori (Sarcopterygii, Oste- notes on other fresh-water limestones. Transactions of the ichthyes) from the Catskill Formation of North Royal Society of Edinburgh, 13, 169–282. America. Journal of Vertebrate Paleontology, 24, 26–40. HUENE, E. von 1943. Ein Onychodus aus dem oberschlesis- DAWSON, J. W. 1868. The geological structure, organic chen Kohlengebirge. Berichte des Reichsamts fu¨r Bodenforschung remains, and mineral resources of Nova Scotia, New Brunswick Berlin, Zweigstelle Wien, 1943, 99–101. 126 PALAEONTOLOGY, VOLUME 49

HUXLEY, T. H. 1862. Note respecting the discovery of a new preserved under the plenary powers in their accustomed usage. and large labyrinthodont (Loxomma allmanni, Huxley) in the Bulletin of Zoological Nomenclature, 27, 97–99. Gilmerton Ironstone. Quarterly Journal of the Geological Society MORRIS, J. 1843. A catalogue of British fossils: comprising the of London, 18, 291–296. genera and species hitherto described; with references to their –––– and ETHERIDGE, R. 1865. A catalogue of the collection of geological distribution and the localities in which they have been fossils in the Museum of Practical Geology, with an explanatory found. Published by the Author, London, 222 pp. introduction. G. E. Eyre. & W. Spottiswoode, London, 381 pp. MURCHISON, R. I. 1839. The Silurian system. J. Murray, JACKSON, J. W. 1952. Catalogue of types and figured specimens London, 768 pp. in the Geological Department of the Manchester Museum. Man- NEWBERRY, J. S. 1856. Description of several new genera chester Museum Publication no. 6, 170 pp. and species of fossil fishes, from the Carboniferous strata of JANVIER, P., LETHIERS, F., MONOD, O. and BALKAS, Ohio. Proceedings of the Academy of Natural Sciences of Phil- O¨ . 1984. Discovery of a vertebrate fauna at the Devonian–Car- adelphia, 8, 96–100. boniferous boundary in SE Turkey (Hakkari Province). Jour- —— 1873. Report of the Geological Survey of Ohio; Vol. 1, Part nal of Petroleum Geology, 7, 147–168. II. Palaeontology. Nevins & Myers, Columbus, USA, 399 pp. –––– and VILLARROEL, C. 2000. Devonian vertebrates from —— 1889. The Palaeozoic fishes of North America. Government Colombia. Palaeontology, 43, 729–763. Printing Office, Washington, USA, 340 pp. JEFFERY, J. E. 1999. The morphology and phylogeny of the —— and WORTHEN, A. H. 1870. Palaeontology of Illinois; Sec- European members of Order Rhizodontida (Pisces: Sarco- tion I, Description of fossil vertebrates. Geological Survey of pterygii). Unpublished PhD thesis, University of Cambridge, Illinois, 4, 343–350. Museum of Zoology, 313 pp. —— —— 1866. Descriptions of new species of vertebrates, —— 2001. Pectoral fins of rhizodontids and the evolution of mainly from the sub-Carboniferous limestone and Coal Meas- pectoral appendages in the tetrapod stem-group. Biological ures of Illinois. 9–134. In WORTHEN, A. H. (ed.). Geologi- Journal of the Linnean Society, 74, 217–236. cal Survey of Illinois, Volume II. Palaeontology. Legislature of —— 2003. Mandibles of rhizodontids: anatomy, function and Illinois, Chicago, USA, 470 pp. evolution within the tetrapod stem-group. Transactions of the NEWMAN, A., M C LEAN, S. G. and HUDSON, D. 1996. A Royal Society of Edinburgh: Earth Sciences, 93, 255–276. catalogue of the type, figured and cited fossil vertebrates in the –––– DAVIS, M. C., DAESCHLER, E. B. and SHUBIN, N. Hancock Museum, Newcastle upon Tyne. J. & P. Bealls Ltd, 2002. Sauripterus Hall, 1843 (Osteichthyes, Sarcopterygii): pro- Newcastle, 161 pp. posed conservation as the correct original spelling [Case 3203]. NIXON, K. C. and WHEELER, Q. D. 1990. An amplification Bulletin of Zoological Nomenclature, 59, 1–5. of the phylogenetic species concept. Cladistics, 6, 211–223. JOHANSON, Z. and AHLBERG, P. E. 1998. A complete prim- OWEN, R. 1840. Odontography, or, a treatise on the comparative itive rhizodont from Australia. Nature, 394, 569–573. anatomy of the teeth; their physiological relations, mode of devel- —— —— 2001. Devonian rhizodontids (Sarcopterygii; Tetrap- opment, and microscopic structure in the vertebrate . odomorpha) from East Gondwana. Transactions of the Royal Hippolyte Bailliere, London, 655 pp. Society of Edinburgh: Earth Sciences, 92, 43–74. PARKER, K., JOHANSON, Z. and WARREN, A. 2005 –––– TURNER, S. and WARREN, A. 2000. First East Gondwa- Strepsodus (Rhizodontida; Sarcopterygii) pectoral elements from nan record of Strepsodus (Sarcopterygii, Rhizodontida) from the Lower Carboniferous Ducabrook Formation, Queensland, the Lower Carboniferous Ducabrook Formation, central Australia. Journal of Vertebrate Paleontology, 25, 46–62. Queensland, Australia. Geodiversitas, 22, 161–169. PORTLOCK, J. E. 1843. Report on the geology of the county of KIRKBY, J. W. and ATTHEY, T. 1864. On some fish remains Londonderry, and of parts of Tyrone and Fermanagh. A. Milliken, from the Durham and Northumberland Coal Measures. Dublin, Ireland, 784 pp. Transactions of the Tyneside Naturalists’ Field Club, 6, 231– RIDE, W. D. L., COGGER, H. G., DUPUIS, C., KRAUS, 235. O., MINELLI, A., THOMPSON, F. C. and TUBBS, P. K. LEIDY, J. 1865. Descriptions of some remains of fishes from 2000. International Code of Zoological Nomenclature. Fourth the Carboniferous and Devonian formations of the United edition. International Trust for Zoological Nomenclature, States. Journal of the Academy of Natural Sciences of Philadel- London, 306 pp. phia, 3, 159–165. ROMER, A. S. 1955. Herpetichthyes, Amphibiodei, Choa- LONG, J. A. 1989. A new rhizodontiform fish from the Early nichthyes or Sarcopterygii? Nature, 176, 126. Carboniferous of Victoria, Australia, with remarks on the phy- —— 1963. The larger embolomerous amphibians of the Ameri- logenetic position of the group. Journal of Vertebrate Paleon- can Carboniferous. Bulletin of the Museum of Comparative tology, 9, 1–17. Zoology, Harvard, 128, 415–454. M‘COY, F. 1854. Contributions to British palaeontology, or first –––– and SMITH, H. S. 1934. American Carboniferous Dipno- descriptions of three hundred and sixty species and several genera ans. Journal of Geology, 42, 700–718. of fossil Radiata, Articulata, Mollusca and Pisces from the Ter- SALTER, J. W. 1861. On the fossils of the South Welsh coal tiary, , Oolitic, and Palaeozoic Strata of Great Britain. fields. 219–296. In: ANON. Iron ores of Great Britain, Part III. MacMillan, Cambridge, 272 pp. Iron ores of South Wales. Memoirs of the Geological Survey of MELVILLE, R. V. and CHINA, W. E. 1970. Opinion 930; Great Britain and of the Museum of Practical Geology. Megalichthys Agassiz, 1835, and Rhizodus Owen, 1840 (Pisces): Longman, Green, Longman & Roberts, London, 296 pp. JEFFERY: CARBONIFEROUS FISH GENERA 127

SCHULTZE, H. P. 1969. Die Faltenza¨hne der rhipidistiiden tions and Annual Report of the North Staffordshire Naturalists’ Crossopterygier, der Tetrapoden und der Actinopterygier-Gat- Field Club, Special Supplement; Addresses and Papers, 184–251. tung Lepisosteus; nebst einer Beschreibung der Zahnstruktur WATERSTON, C. D. 1954. Catalogue of type and figured von Onychodus (struniiformer Crossopterygier). Palaeonto- specimens of fossil fishes and amphibians in the Royal Scottish graphica Italica, New Series, 35, 59–137. Museum, Edinburgh. Transactions of the Edinburgh Geological —— 1970. Folded teeth and the monophyletic origin of tetra- Society, 16, 1–91. pods. American Museum Novitates, 2408, 1–10. WHITE, E. I. 1927. The fish-fauna of the Cementstones of —— 1973. Crossopterygier mit Heterzerker Schwanzflosse aus Foulden, Berwickshire. Transactions of the Royal Society of dem Oberdevon Kanadas, nebst einer Beschreibung von Onych- Edinburgh, 55, 255–286. odontida-Resten aus dem Mittledevon Spaniens und aus dem –––– and BAIRD, D. 1967. Comment on the proposals concern- Karbon der USA. Palaeontographica Abteilung A, 143, 188–208. ing the generic names Megalichthys and Rhizodus (Pisces). —— 1993. Osteichthyes: Sarcopterygii. 657–663. In BENTON, Z.N.(S.) 1690. Bulletin of Zoological Nomenclature, 24, 262. M. J. (ed.). The fossil record 2. Chapman & Hall, London, 845 pp. WHYTE, M. A. 1994. Scottish Carboniferous fresh-water lime- –––– and BOLT, J. R. 1996. The lungfish Tranodis and the tetra- stones in their regional setting. Transactions of the Royal Soci- pod fauna from the Upper Mississippian of North America. ety of Edinburgh: Earth Sciences, 84, 239–248. Special Papers in Palaeontology, 52, 31–54. WILLIAMSON, W. C. 1849. On the microscopic structure of SEDGWICK, A. and M‘COY, F. 1854. A synopsis of the classi- the scales and dermal teeth of some ganoid and placoid fish. fication of the British Palaeozoic rocks, with a systematic descrip- Philosophical Transactions of the Royal Society of London, 1849, tion of the British Palaeozoic fossils in the Geological Museum of 435–474. the University of Cambridge. Cambridge University Press, WOOD, S. P. and ROLFE, W. D. I. 1985. Introduction to the Cambridge, 832 pp. palaeontology of the Dinantian of Foulden, Berwickshire, THOMPSON, J. 1867. On the geology of the Campbeltown Scotland. Transactions of the Royal Society of Edinburgh: Earth District. Transactions of the Geological Society of Glasgow, 2, Sciences, 76, 1–6. 76–89. WOODWARD, A. S. 1890. Vertebrate palaeontology in some THOMSON, K. S. 1966. Megalichthys and Rhizodus (Pisces, American and Canadian museums (part 2). Geological Maga- Rhipidistia): proposal for the stabilization of these generic zine, 7, 455–460. names. Z.N.(S.) 1690. Bulletin of Zoological Nomenclature, 23, —— 1891. Catalogue of the fossil fishes in the British Museum 117. (Natural History), Cromwell Road, S.W., Part 2, containing the TIBERT, N. E. and SCOTT, D. B. 1999. Ostracodes and (Acanthodii), Holocephali, Ichthyodrulites, Os- agglutinated foraminifera as indicators of paleoenvironmental tracodermi, Dipnoi, and Teleostomi (Crossopterygii and Chondr- change in an Early Carboniferous brackish bay, Atlantic Can- ostrean Actinopterygii). Longmans, London, 567 pp. ada. Palaios, 14, 246–260. —— 1902. Preliminary note on a Carboniferous fish fauna from TRAQUAIR, R. H. 1878. On the genus Rhizodus. Proceedings Victoria, Australia. Geological Magazine, 9, 471–473. of the Royal Society of Edinburgh, 9, 657–660. —— 1906. On a Carboniferous fish-fauna from the Mansfield —— 1881. Report on fossil fishes collected by the Geological District, Victoria. Memoirs of the National Museum, Mel- Survey of Scotland in Eskdale and Liddesdale. Part I. – Ganoi- bourne, 1, 1–32. dei. Transactions of the Royal Society of Edinburgh, 30, 15–23. –––– and SHERBORN, C. D. 1890. Catalogue of British fossil —— 1882. Notice of new fish remains from the Blackband Iron- Vertebrata. Dulau, London, 396 pp. stone of Borough Lee, near Edinburgh. Geological Magazine, 9, YOUNG, G. C., LONG, J. A. and RITCHIE, A. 1992. Cross- 540–546. opterygian fishes from the Devonian of : systemat- —— 1890. List of the fossil Dipnoi and Ganoidei of Fife and ics, relationships and biogeographic significance. Records of the the Lothians. Proceedings of the Royal Society of Edinburgh, 17, Australian Museum, Supplement, 14, 1–77. 385–400. YOUNG, J. 1866a. Appendix to report of the committee on —— 1897. Additional notes on the fossil fishes of the Upper the distribution of the vertebrate remains from the North Old Red Sandstone of the Moray Firth Area. Proceedings of the Staffordshire coal field. Reports of the British Association for the Royal Physical Society of Edinburgh, 13, 376–385. Advancement of Science, 1865, 317–320. —— 1903. On the distribution of fossil fish-remains in the —— 1866b. Notice of new genera of Carboniferous Glyptodipte- Carboniferous rocks of the Edinburgh district. Transactions of rines. Quarterly Journal of the Geological Society of London, 22, the Royal Society of Edinburgh, 40, 687–709. 596–608. VOROBYEVA, E. I. and OBRUCHEVA, H. D. 1977. Rhizo- —— 1867. Notice of the occurrence of Rhizodus hibberti in Car- dont crossopterygian fishes (fam. Rhizodontidae) from the boniferous strata in the neighbourhood of Glasgow. Transac- Middle Palaeozoic deposits of the Asiatic Part of the USSR. tions of the Geological Society of Glasgow, 2, 38–39. 89–97. In MENNER, V. V. (ed.). Essays on phylogeny and —— and ARMSTRONG, J. 1871. On the Carboniferous fossils systematics of fossil agnathans and fishes. Nauka, Moscow, 166 of the west of Scotland; their vertical range and distribution with pp. [in Russian]. a general catalogue of the fossils and their mode of occurrence, WARD, J. 1875. On the organic remains of the Coal Measures and an index to the principal localities. Geological Society of North Staffordshire, their range and distributions, with a Press and Transactions of the Geological Society of Glasgow catalogue of the fossils and their mode of occurrence. Transac- [Supplement to Volume 3], Glasgow, 103 pp. APPENDIX 128 AAOTLG,VLM 49 VOLUME PALAEONTOLOGY, Check-list of published rhizodontid species

Taxon and author Type Comments Status

Apedodus priscus Leidy, 1865 A tooth and fragment of an ?opercular, from the Young (1866b) suggested the material resembled Nomen dubium. ‘Old Red Sandstone’ (¼ ?Catskill Formation) from Rhizodus hibberti. Newberry (1889) suggested that Colombia County, Pennsylvania, USA, current the tooth was similar to those of Sauripterus. location uncertain. Archichthys portlocki (Portlock, Teeth BGS GSL 1301, 1302 and 1303 and BGS Type material described and figured by Portlock Valid genus and species. 1843 ex Agassiz, MS) GSM 28500: scales BGS GSL 1304 and BGS GSM (1843) using an Agassiz manuscript name Holop- 28501, from the ?Ivorian (?upper Tournaisian) of tychius portlocki. Agassiz (1844a) later published Londonderry, Northern Ireland. A tetrapod man- the name only (as Holoptychius portlockii) with no dible fragment (BGS GSM 28498) and dermal description or figures. Salter (1861) used the bone (BGS GSM 28502) were also included. name Rhizodus portlocki. The species was trans- ferred to Archichthys by Traquair (1878) because of the similarity of its teeth to those of the type spe- cies, A. sulcidens. Archichthys was synonymized with Strepsodus by Woodward (1891) without explanation. I reject Woodward’s (1891) synony- mization (see main text for details). Archichthys sulcidens Hancock and See Strepsodus sulcidens. Atthey, 1870 Aztecia mahalae Johanson and Right pectoral girdle and humerus, from the Young et al. (1992) described material from the Valid genus and species. Ahlberg, 2001 ?upper of Victoria Land, Antarctica, AMF Antarctic as a rhizodontid species Notorhizodon 113914 (partim AMF 54333, Young et al. 1992). mackelveyi. Johanson and Ahlberg (2001) demon- strated that it was a chimera of several taxa inclu- ding a tristichopterid and a rhizodontid. They used the rhizodontid material to establish Aztecia mahalae. Notorhizodon mackelveyi remains as a valid species of tristichopterid. Barameda decipiens (Woodward, Isolated scale from the Visean of Mansfield, The remains were noted by Woodward (1902) and Valid genus and species. 1906) Victoria, Australia, NMV P10277. later described as Strepsodus decipiens. The species was transferred to Pycnoctenion by Vorobyeva and Obrucheva (1977), and then used by Long (1989) to erect the genus Barameda. Dendroptychius sp. Huxley, in Partial scale, current location uncertain. Type material described by Young (1866b). Wood- Nomen dubium. Huxley and Etheridge 1865 ward and Sherborn (1890) stated that the type was located in the Museum of Practical Geology. How- ever, it is not in the British Geological Survey col- lections, where the MPG material is currently housed. Woodward and Sherborn (1890) also syn- onymised the genus with Strepsodus sauroides. Gooloogongia loomesi Johanson Articulated specimen from the Frasnian of Specimens preserved as natural moulds. Valid genus and species. and Ahlberg, 1998 Canowindra, New South Wales, Australia, AMF 96860a and b. Notorhizodon mackelveyi Young Associated cranial material, from the ?upper See Aztecia mahalae. et al., 1992 Givetian of Victoria Land, Antarctica, CPC 25404. Propycnoctenion nephroides Isolated scale from the Upper Devonian of Based on a single poorly preserved scale. Nomen dubium. Vorobyeva and Obrucheva, 1977 Kzyl-Dzhar Mountain, Kara-Tau mountain range, Kazakhstan, PIN 789 ⁄ 11. Pycnoctenion decipiens See Barameda decipiens. (Woodward, 1906) Pycnoctenion jacuticus Vorobyeva Mandible fragment from the Pennsylvanian of Type specimen is a jaw fragment, somewhat sim- Nomen dubium. and Obrucheva, 1977 Yakutia, Siberia, Russia, PIN 2805 ⁄ 100. ilar to Archichthys portlocki. Pycnoctenion litvinovitschae Isolated scale from the upper Visean of Kypshak Based on a damaged scale, now apparently lost. Nomen dubium. Vorobyeva and Obrucheva, 1977 River, Tselinograd Region, Kazakhstan, GGM 230 ⁄ 21. Pycnoctenion rybinskensis Isolated scale from the Pennsylvanian of Malaya Based on a single scale. Nomen dubium. Vorobyeva and Obrucheva, 1977 Kamala village, Krasnoyarsk Region, Russia, PIN 1741 ⁄ 23. Pycnoctenion siberiacus (Chabakov Isolated scale from the Tournaisian of Kop Village, Described as Strepsodus siberiacus. Used by Nomen dubium. [alt. Khabakov], 1927) Yenisei Province, Siberia, Russia, GRMP 28 ⁄ 2104. Vorobyeva and Obrucheva (1977) as the type spe- cies of their new genus Pycnoctenion. Pycnoctenion tuberculata Isolated scales from the middle Visean of Zhilandy Based on damaged scales. Nomen dubium. Vorobyeva and Obrucheva, 1977 River area, Karaganda Region, Kazakhstan, PIN 2879 ⁄ 3.

Pycnoctenion tuwensis Vorobyeva Isolated scales from the Tournaisian of Sayba Based on isolated scales. Nomen dubium. GENERA FISH CARBONIFEROUS JEFFERY: and Obrucheva, 1977 Village, Tuva, Russia; PIN 1789 ⁄ 2 and 2a. Rhizodus angustus Newberry, 1856 Lower jaw from the ?Pennsylvanian of Ohio, USA, Specific distinction based on slenderness of teeth. Nomen dubium. current location uncertain. Rhizodus anceps Newberry, 1889 Mandible from the Visean of Alton, Illinois, USA, The holotype, from the St Louis Limestone Forma- Junior synonym of Rhizodus AMNH 224. tion, is indistinguishable from R. hibberti. hibberti. Rhizodus ferox Morris, 1843 None. Given incorrectly as the species name suggested by Nomen dubium. (in errore) Owen (1840) for his new genus Rhizodus. Owen in fact used a version of Agassiz’s manuscript name, Rhizodus hibbertii. Rhizodus garneri (Murchison, Lost, according to Ward (1875). Published as Holoptychius garneri Agassiz, 1844a. Nomen dubium. 1839 ex Agassiz, MS) The species was transferred to Rhizodus by Ward (1875), although he noted the type material was lost. Woodward (1891) suggested it was a junior synonym of Strepsodus sauroides. Rhizodus gracilis M‘Coy, in Dentary from the Brigantian of Gilmerton, Synonymized with R. hibberti by Traquair (1878) Junior synonym of Rhizodus Sedgwick and M‘Coy 1854 Edinburgh, Scotland, SME 4713. and Woodward (1891). hibberti. 129 Appendix. Continued 130

Taxon and author Type Comments Status 49 VOLUME PALAEONTOLOGY,

Rhizodus granulatus (Agassiz, Agassiz (1844a) did not give the details of any Holoptychius granulatus Agassiz, 1844a (name only) Junior synonym of Rhizodopsis 1844a) specimens, but gave the type locality as Manches- transferred to Rhizodus by Salter (1861). Trans- sauroides. ter. ferred to Rhizodopsis by Barkas (1873) and to Rhi- zodopsis sauroides by Woodward and Sherborn (1890). Rhizodus hardingi Dawson, 1868 See Strepsodus hardingi. Rhizodus hibberti Owen, 1840 Dentary tusk from the Asbian of Burdiehouse, Type material described by Hibbert (1835) as Valid genus and species. (by ICZN ruling) Edinburgh, Scotland, NMS GY 1950.63. Megalichthys hibberti using a manuscript name from Agassiz. Owen (1840) erected the genus Rhiz- odus, retaining the specific name (there spelled hib- bertii). See Thomson (1966), White and Baird (1967), Melville and China (1970) and Andrews (1982, 1985) for more details. Rhizodus incurvus Newberry, 1856 Cranial bones and teeth from the ?Pennsylvanian Specific distinction based on tooth shape, com- Nomen dubium. of Ohio, USA, current location uncertain. pared with Rhizodus gracilis, not the type species Rhizodus hibberti. Rhizodus lanciformis Kirkby and None. Used by Kirkby and Atthey (1864), probably a Nomen dubium. Atthey, 1864 (in errore) misspelling of R. lancifer. Woodward and Sherborn (1890) stated that the teeth figured by Kirkby and Atthey (1864) belong to the tetrapod Loxoma all- mani. Rhizodus lancifer Newberry, 1856 Palatal tusk from the ?Pennsylvanian of Ohio, Based on an isolated tooth from Linton, Ohio. Valid species of tetrapod genus USA, OSU 4500. Used as the type material for the tetrapod species Anthracosaurus. Anthracosaurus lancifer by Romer (1963). Rhizodus occidentalis Newberry Isolated scale from the ?Myachkovskian of Morris, Transferred to the dipnoan genus Sagenodus by Synonym of lungfish species and Worthen, 1866 Illinois, USA, current location uncertain. Cope (1897). Sagenodus [formerly Rhizodus] retic- Ctenodus cristatus. ulatus was synonymized with S. occidentalis by Ro- mer and Smith (1934). Baird (1978) and Bardock (1979) considered S. occidentalis to be a junior synonym of the dipnoan Ctenodus cristatus. Rhizodus ornatus Traquair, 1878 See Screbinodus ornatus. Rhizodus portlocki (Portlock, 1843 See Archichthys portlocki. ex Agassiz, MS) Rhizodus quadratus Newberry, Large, thin scale from the Myachkovskian Cannel Newberry (1873) noted that they may be the scales Nomen dubium. 1873 Coal of Linton, Ohio, USA, current location of Rhizodus lancifer. Other material also mentioned uncertain. in Newberry (1889). Rhizodus reticulatus Newberry and Isolated scale from the Myachkovskian of Mazon Newberry (1873) suggested that the scales may See Rhizodus occidentalis. Worthen, 1870 Creek, Illinois, USA, YPM 3268. belong to Rhizodus lancifer. Woodward (1891) transferred the species to the lungfish genus Sagen- odus. Rhizodus sauroides (Binney, 1841) See Strepsodus sauroides. Rhizodus striatus (Agassiz, 1844a) Unknown. Described as Holoptychius striatus Agassiz, 1844a Nomen dubium. (name only). Moved to Rhizodus by Etheridge (1888), although no reasons were given and no material was figured. Sauripteris taylori Hall, 1843 See Sauripterus taylori. Sauripterus anglicus Woodward, Scales from the Upper Devonian of Farlow, Shrop- Erected by Woodward based on a few scales and a Valid species of porolepiform 1891 shire, England, NHM P200 and P201. tooth. Used by Ball et al. (1961) as the type species Pseudosauripterus anglicus of a porolepiform genus Pseudosauripterus. Sauripterus crassidens Traquair, Tooth from the Upper Devonian of Alves, Moray, Species assigned to Sauripterus due to similarity of Nomen dubium. 1897 Scotland, NMS GY 1936.38.12. teeth to Sauripterus favosus, not to the type species, S. taylori. Sauripterus favosus (Agassiz, Lost by Agassiz, according to Woodward (1891). Type material of Glyptosteus favosus Agassiz, 1844a, Nomen dubium. 1844a) p. 179 (name only) was later figured by Agassiz (1844b, p. 61) as Bothriolepis favosa and then lost. Woodward (1891) used the figures to assign some new material to the species and to transfer it to Sauripterus. However, the material Woodward assigned to Sauripterus favosus is scrappy and can- not be compared with the type species of Sauripte-

rus, S. taylori. GENERA FISH CARBONIFEROUS JEFFERY: Sauripterus taylori Hall, 1843 Pectoral fin and girdle from the Famennian of Described by Hall (1840) as Sauritolepis taylori Valid genus and species. Blossberg, Pennsylvania, USA, part of AMNH before replacing that name with Sauripteris [sic] 3341. taylori. See Jeffery et al. (2002) for details on the type series and spelling of Sauripterus. Sauritolepis taylori Hall, 1840 See Sauripterus taylori. Screbinodus ornatus Articulated anterior trunk from the Brigantian of Traquair (1878) erected this as a new species of Valid genus and species. (Traquair, 1878) Gilmerton, Edinburgh, Scotland, NMS GY 1878.4. Rhizodus. Andrews (1985) made it the type species of Screbinodus. ?Strepsodus anculonamensis Complete specimen from the Ivorian (Tn3) of Based on a complete, articulated specimen, Junior synonym of Strepsodus Andrews, 1985 Foulden, Berwickshire, Scotland, NMS GY although the taxonomic problems of Strepsodus sauroides. 1980.40.36. (see main text) prevented a more certain as sign- ment to that genus. Strepsodus arenosus Hay, 1900 Isolated scale from the base of the Mississippian at The holotype is probably a rhizodontid scale, but Nomen dubium. Collier’s Station, Blair county, USA, USNM 4855. shows no reliable taxonomic characters. 131 Appendix. Continued 132

Taxon and author Type Comments Status 49 VOLUME PALAEONTOLOGY,

Strepsodus brockbanki Davis, 1891 Disarticulated bones and teeth from Moscovian Based on a poorly preserved specimen, which Nomen dubium. (Westphalian B) of Levenshulme, Manchester, shows no definite rhizodontid characters. England, MM LL222. Strepsodus dawsoni Hay, 1900 Single scale from the Podolskian or lower Based on a single scale originally figured by Daw- Nomen dubium. Myachkovskian of Pictou, Nova Scotia, Canada, son (1868) as Rhizodus lancifer. Hay (1900) distin- RM 3076. guished it from the holotype of Strepsodus hardingi (also from Nova Scotia, Canada) by its younger age. Strepsodus decipiens Woodward, see Barameda decipiens. 1906 Strepsodus hardingi (Dawson, Fragment of mandible from the Ivorian of Horton Originally described as Rhizodus hardingi. Moved Currently under study by Mr M. 1868) Bluff, Nova Scotia, Canada, RM 2706. to genus Strepsodus by Woodward (1891). Brazeau, Redpath Museum Strepsodus minor Traquair, 1890 Scales and a tooth from the Tournaisian of Pitcort- Little material attributed besides the type series. Junior synonym of Strepsodus hie, Fife, Scotland, NMS GY 1881.43.42. sauroides. Strepsodus portlocki See Archichthys portlocki. (Portlock, 1843 ex Agassiz, MS) Strepsodus sauroides A single tooth from the Moscovian (Westphalian Published as Holoptychius sauroides. Young (1867) Valid genus and species. (Binney, 1841; pars.) B) Pendleton coalfield, current location uncertain. used the name Rhizodus sauroides. See main text Neotype SME 4721 (Text-fig. 4). for further details of the taxonomy of S. sauroides. Strepsodus siberiacus Chabakov See Pycnoctenion siberiacus. [alt. Khabakov], 1927

Strepsodus striatulus Traquair, Seven teeth from the Pendleian of Burghlee, Edin- Traquair used the spelling ‘Borough Lee’ for the Junior synonym of Strepsodus 1882 burgh, Scotland. Listed as NMS GY 1950.38.73 by location. Specimen from Traquair’s collection sauroides. Waterston (1954), but now apparently lost. (SME 4740) apparently identical to Strepsodus sauroides. Strepsodus sulcidens (Hancock Left mandible, NEWHM G59.64, from the Type material first described as Rhizodus hibberti Junior synonym of Archichthys and Atthey, 1870) Kashirskian (Westphalian B) of Newsham, (Hancock and Atthey 1868), but later used to portlocki. Northumberland, UK establish Archichthys sulcidens. Genus Archichthys was synonymised with Strepsodus by Woodward (1891) without explanation. I reject Woodward’s (1891) synonymization, although A. sulcidens is a junior synonym of A. portlocki (see main text for details).