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Revised Taxonomy of Amphibians Albanerpetontid

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Revised Taxonomy of Amphibians Albanerpetontid Revised taxonomy of albanerpetontid amphibians JAMES D. GARDNER Gardner, J.D. 2000. Revised taxonomy of albanerpetontid amphibians. - Acta palaeon- tologica Polonica 45, 1,55--70. Characters of thejaws and frontals are often used to differentiate albanerpetontid genera and species, yet the reliability ofthese characters has rarely been examined. Frontals are diagnostic for the genefaAlbanerpeton and Celtedens and for species in the latter genus. The value of frontals at the specific level in ce ltedens may be inflated by lack of informa- tion about variation injaw structure. characters of the frontals, jaws, and body size dif- ferentiate speciesof Albanerpeton. Differential diagnoses are presented for the Albaner- petontidae based on cranial and vertebral characters and for the two named genera based on frontal characters. Each genus is characterized by one autapomorphy: fused frontals triangular inAlbanerpeton and internasal process bulbous in celtedens. An enigmatic albanerpetontid from the Middle Jurassic (upper Bathonian) of England has a unique mixture of frontal and premaxillary character states that precludes it from being included in either Celtedens or Albanerpeton.Thisleaves the oldest occurrences of the two genera in, respectively, the Late Jurassic (Kimmeridgian) and Early cretaceous (latest Aptian/ earliest Albian). Key words: Albanerpeton, Albanerpetontidae, Allocaudata, Amphibia, Celtedens, Lissamphibia, taxonomy. James D. Gardner [[email protected]], Laboratory for vertebrate paleontology and Deparxnent ofBiological sciences, university ofAlberta, Edmonton, AtbertaT6G 289 Canada. Introduction The AlbanerpetontidaeFox & Naylor, 1982 ne salamander-like,Middle Jurassicto Miocene amphibians that are of general interest becausethey constitute a clade of lissamphibiansdistinct from caudates,salientians, and gymnophionans.Two genera, containing at least nine species,are currently recognized:the type genusAlbanerpeton Estes& Hoffstetter,I9T6,from the Lower Cretaceous-Paleoceneof the Norttr American westernInterior andMiocene of France@stes & Hoffstetter1976; Estes 19g1; Fox & Naylor 1982;Gardner r999a, b, c) and celtedensMcGowan & Evans.1995 from the 56 Revised taxonomy of albanerpetontid amphibians; GARDNER Lower Cretaceous,Upper Jurassic,and, purportedly, Middle Jurassicof Europe (I\4cGowan& Evans1995; McGowan1996,1998; McGowan & Ensom1997). Jaws and frontals are the most commonly recoveredalbanerpetontid elements at most sites,and charactersof theseelements are widely usedto diagnosegenera and species(Estes 1981; Nessov 1981; Fox & Naylor 1982;McGowan & Evans 1995; McGowan 1998;Gardner l999a,b, c; Rage& Hossiniin press).In many cases,the re- liability of such charactershas not been adequatelyestablished. This is the result of a combinationof factors:scarcity of well preservedspecimens; a reluctanceon the part of someworkers to examineappropriate comparative collections; and misinterpreta- tions of structure.To cite a recentexample, McGowan (1998) used only frontal charac- ters to diagnoseAlbanerpeton, Celtedens, and their respectivespecies. McGowan's (1993)claims aboutthe diagnosticutility of frontals for Albanerpetonand its species are suspectbecause he consideredjusttwo ofthe four namedspecies then includedin the genus(I recognizeseven species), examined only material of the type speciesA. inexpectatumEstes& Hoffstetter,1976, andmisinterpretedthe structureof the frontal inA. galaktionFox & Naylor, 1982based on photographs(Fox & Naylor I982:fig.ld, e) of an incompletespecimen. McGowan's (1998)reliance on frontal charactersalone to diagnosespecies of Albanerpetonis alsoat oddswith observationsthat charactersof the jaws are also useful for this purpose(Estes 1981; Fox & Naylor 1982; Gardner I999a,b, c; Rage& Hossiniin press). Here I assessthe taxonomicvalue of charactersfor albanerpetontids,with empha- sis on frontals and jaws. Isolated examplesof theseare availablefor all speciesof Albanerpeton(Estes & Hoffstetter1976;Fox & Naylor 1982;Gardnet I999a,b, lun- published data), except for a new Turonian speciesknown only by its distinctive holotype premaxilla (Gardner I999c). Frontals are also reasonablywell known for Celtedens,bothas disassociatedelements from indeterminatespecies and articulated in the respectiveholotype skeletonsof both namedspecies, C. megacephalus(Costa, 1864)and C. ibericusMcGowan & Evans,1995 (McGowan & Evans1995; McGo- wan 1996,1998;McGowan & Ensom 1997).Jaws in theseskeletons are difficult to studybecause they arepreserved in articulationand, in the caseof the holotypeof C. megacephallzs,also poorly preserved.The only isolated,reasonably well preserved jaws availablefor Celtedensare incomplete premaxillae and dentaries of an indetermi- nate speciesfrom the Lower Cretaceousof Purbeck,England (McGowan & Ensom 1997).In the secondpart of this paper,I provide reviseddiagnoses and selectcom- ments for the Albanerpetontidae and its genera,then examine the identity of a pur- portedCeltedens-llke taxon (McGowan 1996,1998)from the Middle Jurassicof Eng- land. Osteologicalterms and measurements follow Gardner(1999a), with oneexception. 'lacrimal For the premaxilla, I proposethe replacementterm notch' for the indentation alongthe laterodorsaledge ofthe parsdorsalis - this notchreceived the largelacrimal, not the reducedprefrontal (Gardnerin press).I follow McGowan & Evans (1995) in placingalbanerpetontids within the LissamphibiaHaeckel, 1866 and Milner (1988)in regardinglissamphibians as crown-cladetemnospondyls. Institutional abbreviations: BMNH, Natural History Museum, London, England; DORCM, Dorset County Museum, Dorchester,England; FMNH, Field Museum of NaturalHistory, Chicago, USA; LH, Las Hoyascollection, Museo de Cuenca,Cuenca, ACTA PALAEONTOLOGICAPOLONICA (45) (I) 57 Spain; MNHN.LGA, La Grive-St. Alban collection, Mus6um National d'Histoire Naturelle,Paris, France; MNP, Museodi Paleontologiadell'Universitd di Napoli, Na- ples, Italy; RTMP, Royal Tyrrell Museum of Palaeontology,Drumheller, Canada; UALVP, University of Alberta Laboratory for Vertebrate Paleontology, Edmonton, Canada;UCK, University CollegeLondon, London, England; and UCMP, University of California Museumof Paleontology,Berkeley, USA. Taxonomic characters for albanerpetontids Frontals (Figs 1, 2;Table 1). - Albanerpetontidfrontals are solidly fusedmedially and have a prominent internasalprocess, two pairs of slots anterior to the orbital mar- gin for mortise and tenon style articulation with posterior endsof the paired nasalsand prefrontals, prominent ventrolateral crests adjacent to the orbital margin, and dorsal omamentcomprised of polygonal pits enclosedby ridges. Comparisonswith other temnospondylsindicate that the frst three of thesecharacter states are synapomorphic for the family (Gardnerunpublished data). Table 1. Frontal characters for albanerpetontid genera and species. References are: lMcGowan & Ev- ans1995;2McGowan 1998;3Gardnerl999a;aRage &Hossiniinpress;5S.E. Evans personalcommu- nication 1999. l. Dorsal or ventral outline of frontalsl,2'3. 2. Midline length of frontals relative to width across posterior edge between lateral edges of ventrolateralcrests2'3'4. 3. Dorsal or ventral outline of internasal processl,2'3. 4. Length of internasal processrelative to width acrossbase2. 5. Pattem of contact between internasal process and paired nasals3. 6. Form of anterolateral processs. 7. Medial emargination of posterior slot for receipt of prefrontal2. 8. Width of frontals between slots for receipt ofprefrontal relative to width across posterior edge of bone2. 9. Angle at which lateral wall of frontal diverges posterolaterally from midline2. 10. Position of anterior limit of orbital margin relative to midline length of frontals2. 11. Dorsal or ventral outline of orbital marginl'2. | 2. Form of ventrolateral crests. 13. Pattern of dorsal omament2. Table 1 lists the 13 frontal charactersthat have been used to differentiateAlbaner- peton,Celtedens, and their respectivespecies. Seven characters (1-3, 6,'7,10, and 11) differ betweenAlbanerpeton and Celtedens, as follows (Figs | , 2): l, outlineof frontals triangularin Albanerpelo/z,versus more nearlyhourglass- or bell-shapedin Celtedens; 2, midline length of frontals relative to width acrossposterior edge between lateral edges of ventrolateralcrests less than about l.2inAlbanerpeton, versus more than about 1.2 in Celtedens;3,outline of intemasalprocess triangular inAlbanerpeton,versus bulbous in Celtedens;6,anterolateral process distinct from main body offrontal, extendingantero- laterally, and pointed distally tn Albanerpelor2,versus a poorly differentiated,broadly rounded shoulder in Celtedens;7, dorsal and ventral edges of slot for receipt of prefrontal excavated medially in Albanerpeton, yetslrs unexcavated tn Celtedens; 58 Revised taxonomy of albanerpetontid amphibiarcs.'GARDNER 10, anterior end of orbital margin, as marked by posterior edge of slot for receipt of prefrontal, lies approximatelyin line with, or posterior to, anteroposteriormidpoint of frontals inAlbanerpeton, vetsusanterior end of orbital margin lies anterior to midpoint of frontals in Celtedens;and 11, outline of orbital margin uniformly shallowly curvedto nearly straight in Albanerpelonoversus margin more concavemedially and, occasion- ally, deflectedposterolaterally near posterior end in Celtedens.Where known, the pat- tern of frontal-nasalcontact (character5) also differs betweenthe two genera.Ventral facetson the internasalprocess of an incompletefrontal (Fig. 2A) of Celtedenssp. from Purbeck indicate
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