ANTECHINUS PUTEUS (MARSUPIALIA: DASYURIDAE), A NEW FOSSIL SPECIES FROM THE TEXAS CAVES, SOUTHEASTERN QUEENSLAND

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1 ANTECHINUS PUTEUS (MARSUPIALIA: DASYURIDAE), A NEW FOSSIL SPECIES FROM THE TEXAS CAVES, SOUTHEASTERN QUEENSLAND Stephen Van Dyck Van Dyck, S., (Marsupialia: Das>uridae>, a new fossil species from the Texas Caves, southeastern Queensland. Aust. Mammal. 5: 59*68. A ntechinus puteux, a new fossil species of Anitchinus is described from a Pleistocene deposit in the Texas Caves of southeastern Queensland. This species most closely resembles A. flavipes (Waterhouse) but differs from it in possessing a single-rooted, obliquely oriented Pn9and reduced talonids on Ma and M>. Stephen Van Dyck. Queensland Museum. Gregory Terrace. Fortitude Valley. /in\bartc. Queensland. Australia Manuscript received 5 January, IN 1973 AND 1975 M. Archer and coworkers recovered fossil remains from the sediments of Russenden Cave, part of a complex known as the Texas Caves, on Viator Hill southeastern Queensland (Archer 1978). These caves have since been flooded by the Glenlyon Dam, constructed under the authority of the Dumarcsq-Barwon Rivers Commission. Russenden Cave's fossilifcrous sediments are mainly detrital accumulations from solution pipes and the Russenden fauna is regarded to be Pleistocene in age (Archer 1978). In 1978, Archer summarized the fossil faunas from the Glenlyon area and drew particular attention to a left dentary (F8445) of a species of Aniechinus found in the 1975 Clean Up (CUV level of the Bone Chamber excavation, Russenden Cave (VR-14). He noted that the specimen was longer than any modern specimen of Aniechinus flavipes in such features as the length Ms-Mi, M3 length and width, and width of M2. It had a short premolar row and bulbous crown bases. Archer was satisfied that this antcchinus could not be referred to any known species unless to A. flavipes. Subsequent X-ray photography and excavation of the encrusted alveolus of P:l revealed, during the present study, that F8445 also had a single-rooted P3. A recent search among the other dasyurid dentaries collected by Archer and co-workers from the Texas Caves resulted in the discovery of another similar specimen of a species of Aniechinus, F8038, a right dentary collected from the Rear Entrance of Russenden Cave in February, Although this specimen is relatively more worn than F8445, it nevertheless retains an intact, single-rooted P3. It is clear that this species of Aniechinus is distinct. It is described below as Antechinus puteus. Dental terminology follows that used by Archer (1976) and Van Dyck (1982) ani tooth number follows the nomcnclaturai system established by Archer (1978). The way in which dental measurements have been made is shown in Fig. I. Generic reallocations of the species regarded by Ride (1970) to be in Aniechinus follows Archer (1982). SYSTEMATICS Antcchinus puteus n. sp. (Figs 2-4b) Hololype: QM F8038 (Fig. 2): Right dentary fragment of an adult with Pa and collected by M. Archer and coworkers, February, Type Locality: Rear Entrance. Russenden Cave, Viator Hill, Glenlyon (28 51*S 15! 28,E), southeastern Queensland.

2 60 AUSTRALIAN MAMMALOGY Paratype: Q M F8445 (Figs 3-4b); Left dentary fragm ent with P2, youn g adult, co lle cted from the B one Cham ber, Russenden C ave by M. A rch er and co-workers. February Diagnosis'. A m ech in u s puieus is a large sp ecies o f A m ech in u s m ost closely resem b ling A. flavipes put differing from it in having an obliquely oriented, single-rooted P.^, a greater bu ccal orientation o f the talonid en d oloph s in M 2-M 4 and a m ore reduced paracon id on M a. T h e above features, com b in ed with the m uch w ider m olar width and short prcm olar row. separate it from the follow in g species: A. godm ani, A. stuartii. A. swainsonii. A. bell us, A. m inim us, A. m elanurus, A. naso. A. w ilhelm ina and A. mayeri. A. puteus differs from Par ant echin us apicalis in having a sh orter and w ider P2, a taller, m ore robust P.t, a less redu ced paraconid on M a, a m uch larger m ctacon id on M 2 and a red u ced talonid on M.-.. It differs from P seudan tech in us m acdon nelien sis and Dasykaluta rosa m on d ae in bein g m uch larger, in possessin g a P ;{, a less reduced paraconid on M 〇, m uch reduced talonids on M s and and high en tocon id s on M 3 and M,. Il differs from Parantechinus bilarni in having a w ider P2, a taller and m ore robust P :,, a less reduced paraconid on M 2, a m uch larger m cta con id on M 2 and less reduced talonids on M :, and M 4. Etym ology: **puteus,t is Latin for **deep wclp\ It is a double allusion; first to the collectin g site, the now flooded caverns of the T exas Caves, and second to a proverb o f H eraclitus 一 ^truth lies at the bottom o f a well,where the rest o f this Antcchinus remains. D escription: Pa (Fig. 2c) is the only prem olar present in the dcnlary o f F8038. It has a single, broad, peg-like root. The tooth is elliptical in outline. Its prominent primary cusp lies anterior to the centre of the tooth. A ridge leads anteriorly from its apex to its base where it joins the weak, continuous buccal and lingual cingula. Posteriorly, a worn crest descends to the cingular shelf and cuspulc. A ridge descends buccally from the apex o f the primary cusp to the buccal cingulum. The P;, is aligned obliquely with respect to the lon g axis o f the tooth row, and with its buccally oriented primary cusp at an angle o f approxim ately 20o toward the anterior from the transverse plane. In this position, alm ost the entire buccal cingulum o f P:, lies in close juxtaposition with the indistinct anterior cingulum o f M2. The Pa of A. puteus is com pared in Figs 2a,c with the P:, o f A. flavipes rubeculus (Fig. 4c). T he M2 (Fig. 2c) talonid is wider than the trigonid. The anterior cingulum is poorly developed. The parastylid corner of

3 VAN DYCK: ANT ECHINUS PUTEUS, AN EXTINCT DASYURID 61 H k- 2. Antechmus puieu\. holotypc QM F8038. a. Lateral vicu of P, M.. b. Occlusal I view of P M.. c. Lateral view of P. and M ; (junction obscured by glue). <1. Lateral view of M :. c. Lateral view of M«. f. Occlusal view of Mt. is ihc most anterior portion of the crown. A well-developed posterior cingulum continues into the buccal cingulum which terminates slightly posterior of the longitudinal midline through the protoconid. There is no lingual cingulum. The paraconid is reduced and slightly higher lhan the crown height of the worn entoconid. The protoconid is the tallest cusp of the trigonid. The mclaconid is worn but unreduced and shorter lhan the protoconid. The hypoconid is just shorter than the mctaconid. and the hypoconid is taller than the cntoconid. The entoconid is worn, reduced, and subcqual in height to the paraconid. The paracrislid is almost vertical from the protoconid to the paracristid fissure, and horizontal between the paracristid fissure and ihc paraconid. The metacristid inclines more steeply from the protoconid to the mctacristid fissure than from the mctaconid to the mctacristid fissure. The metacristid and hypocristid arc approximately oblique to the long axis

4 62 AUSTRALIAN MAMMALOGY of the dentary. The cristid obliqua extends from the hypoconid to the posterior wall of the trigonid. intersecting the trigonid at a point below the protoconid, and buccal to the longitudinal mid-line through the protoconid. The hypocristid extends from the hypoconid to the hypoconulid without verging towards the entoconid. From the base of the metaconid. ihe talonid endolph takes a sharp buccal orientation. The M.i (Fig. 2d) trigonid width is the same as that of the talonid. The anterior cingulum is well-developed in comparison with that of M2 and terminates lingually to accommodate the hypoconulid notch. A prominent posterior cingulum continues into the buccal cingulum, less prominently al the base of ihe hypoconid and terminates slightly anterior to the cristid obliqua trigonid junction above. The worn paraconid is reduced and is the smallest trigonid cusp. It is shorter in height than ihe metaconid. The metaconid is taller than the hypoconid and is slightly taller than the badly worn protoconid. The hypoconid is

5 VAN DYCK: ANT ECHINUS PUT BUS. AN EXTINCT DASYURID 63 Fig. 4. a. Antechinu.\ puteus paratype, lateral view of Ms. b. A. puieus paralypc. occlusal view of M«. c. A. flavipe% rubeculus, lateral view of Pn and Ms. d. A. flavipes rubeculus, (Kclusal view of Mn and M.. subcqual in height to the cntoconid. The worn entoconid is very small, peg-like and elliptical in crown view, not connected to the hypoconulid by a crest but connected to the metaconid by a low cresl. The hypoconulid is slightly higher than the worn entoconid. The paracristid is inclined from the worn protoconid to the paracristid fissure and inclined from the fissure to the paraconid. The mctacristid is equally inclined from the worn protoconid to the metacristid fissure as from the metaconid to the metacristid fissure. The cristid obliqua extends to the base of the protoconid, intersecting the trigonid at a point more lingual to the point directly below the worn protoconid tip and well buccal to the metacristid fissure. On the Ms there is no trace of a lingual cingulum. From the base of the metaconid to the tip of the hypoconulid the talonid endoloph takes a sharp buccal orientation. The M4 (Figs 2c, f) trigonid is wider than the talonid. The anterior cingulum is as in the Ma. The posterior cingulum is comparable in length to the anterior cingulum and it docs not continue into the buccal cingulum. The buccal cingulum is confined to the area between the base of the protoconid and the hypoconid as a thickened bulge of enamel. There is no trace of a lingual cingulum. The paraconid is much reduced and is the smallest trigonid cusp. It is shorter than ihc hypoconid and is subequal in height to the hypoconulid. The small entoconid is not connected to the hypoconulid or the metaconid by a crest. The hypoconulid is higher than ihc entoconid. The cristid obliqua intersects the trigonid at a point more lingual to the longitudinal mid-line through the protoconid tip than From the base of the metaconid. to the tip of the hypoconulid, the talonid endoloph takes a sharp buccal orientation. Meristic changes along the tooth-row as follows. The paraconid height increases from M2 to M.i but is reduced in M,. The

6 64 AUSTRALIAN MAMMALOGY melaconid length at the base of the cusp decreases from The cntoconid of Ma is largest with the M4 entoconid larger than that on M2. The hypoconulids of are subcqual in height. The protoconids of M2 and M, arc subequal in height and both arc taller than the worn (and damaged) protoconid of M3. The hypoconids decrease in size from The talonid of M ( is narrower than the trigonid, the talonid of M3 i:»equal to the trigonid width, the talonid of Ma is greater than (he trigonid width. The paracristid of is shorter than that in Ma which is shorter than that crest in M». The metacristid length increases from The cristid obliqua intersects the trigonid in a progressively more lingual position from M- to M4. The hypocristid of M.t is greater in length than the subequal condition in M2 and M4. The anterior cingulum decreases in length from The posterior cingulum is longest in decreasing in M_. and is shortest in M<. The buccal cingulum decreases in length from M 〇 to M*. Description of paratype QM F8445: The P2 (Fig. 3c) is the only premolar present on the dentary of F8445. Il is broad and has a semi-triangular outline in crown view. Its primary cusp lies anterior to the centre of the tooth. A ridge leads anteriorly from its apex to its base where it joins the continuous buccal and lingual cingula. Posteriorly a worn crest descends to the cingular shelf and cuspule. The P3 (Figs 3c,d) alveolus is narrow, elliptical and in close juxtaposition with the posterior root of P2 and the anterior root of M... It is transversely oriented lo the long axis of the tooth-row. The alveolus indicates the presence of a single root for P.i. The crowded, more specialized condition of the prcmolar row is not a juvenile condition in P8445. Although not an old adult there is sufficient wear, particularly on M:t to compare with adult specimens of A. flavipes flavipes and A. f. rubeculus. X-ray photography of F8445 showed the absence of a deciduous first molar Mj. The M2 (Fig. 3d) talonid is wider than the trigonid. The anterior cingulum is poorly developed. The parastylid corner of M2 is the most anterior portion of the crown. The well-developed posterior cingulum continues into the buccal cingulum which terminates slightly posterior of the longitudinal midlinc through the protoconid. There is no lingual cingulum. The paraconid is reduced to a height subequal to the crown height of the entoconid. The protoconid is ihc tallest cusp of the trigonid. The metaconid is unreduced and shorter than the protoconid. The hypoconid is just shorter than the metaconid and taller than the entoconid. The entoconid is reduced and slightly taller than the paraconid. The paracristid is indistinct and almost vertical from the protoconid to the paracristid fissure and horizontal between the paracristid fissure and the paraconid. The metacristid is more steeply inclined from the protoconid to the metacrislid fissure than from the metaconid to the metacristid fissure. The metacristid and hypocristid arc approximately transverse lo the long axis of Ihc dentary. The cristid obliqua extends from the hypoconid to the posterior wall of the trigonid. intersecting the trigonid at a point below the protoconid and slightly buccal to the longitudinal mid-line of the protoconid. The hypocristid extends from the hypoconid to the hypoconulid without verging toward the entoconid. From the base of the entoconid, the talonid endoloph takes a sharp buccal orientation. The hypoconulid is high, almost reaching the level of the paracristid and contacting just lingual lo the midpoint of the paracristid (between the paraconid and the paracristid fissure). The Ms (Fig. 3c) trigonid is wider than the talonid. The anterior cingulum is welldeveloped compared to M..t and terminates lingually to accommodate the hypoconulid notch. The prominent posterior cingulum continues into the buccal cingulum, less prominently at the base of of hypoconid

7 VAN DYCK: AWTTC 謂 US PUm/S, AN EXTINCT DASYURID 65 and terminating slightly anterior to the cristid obliqua 一 trigonid junction above. The paraconid is reduced and is the smallest trigonid cusp, subequal in height to the metaconid. The metaconid is taller than the hypoconid and much shorter than the protoconid. The hypoconid is taller than the entoconid. The cntoconid is small, peglike and almost round in crown view, not connected to the hypoconulid by a crest but connected to the metaconid by a low crest. The hypoconulid is lower than the cntoconid and contacts M4 just lingual to the midpoint of the paracristid (between the paraconid and the paracristid fissure) and level with the paracristid fissure. The paracristid is vertical from the protoconid to the paracristid fissure and inclined from the fissure to the paraconid. The metacristid is more steeply inclined from the protoconid to the metacristid fissure, than from the metaconid to the metacristid fissure. The cristid obliqua extends to the base of the protoconid intersecting the trigonid at a point directly below the protoconid tip and well buccal to the metacristid fissure. On M.-, there is a trace of a lingual cingulum as two thickened bulges of enamel, one directly between the paraconid and the metaconid, the other directly between the metaconid and the entoconid and below the metaconid 一 entoconid crest. The rest of the morphology of Ms is as in M The M4 (Figs. 3f, 4b) trigonid is wider than the talonid. The anterior cingulum is as in M-t. The posterior cingulum is comparable in length to the anterior cingulum and docs not continue into the buccal cingulum. The buccal cingulum is confined to the area between the base of the protoconid and the hypoconid as a thickened bulge of enamel, less well-developed than the anterior and posterior cingula. There is a trace of a lingual cingulum as in M:{ but less clearly defined. The paraconid is the smallest trigonid cusp and is taller than any talonid cusp. The entoconid is not connected to the hypoconulid or the metaconid by a crest. The hypoconulid is subequal to the entoconid and contacts M:, midway along the paracristid (between the paraconid and the paracristid fissure). The hypoconulid tip is level with the paracristid fissure of Mr>. The cristid obliqua intersects the trigonid at a point more lingual to the longitudinal mid-line through the protoconid tip than in Ma. The rest of the morphology is as in M.v The Mn (Fig. 4a) trigonid is wider than the talonid. The anterior cingulum is as in M3. The posterior cingulum is absent. The buccal cingulum is confined to the area between the trigonid and the base of the hypoconid below its tip. The paraconid is just shorter than the metaconid. The talonid has one main cusp (the hypoconid) and has the cntoconid and hypoconulid suppressed. The hypoconid is reduced relative to M4. The cristid obliqua is a high crest which intersects the trigonid below and lingual to the metacristid fissure, this being more markedly lingual than the intersections of the cristid obliqua for M2 一 M4. At the point immediately before intersection with the trigonid a swelling of enamel on the cristid obliqua forms a minute cusp approximately half the height of the hypoconid. Meristic changes along the tooth-row are as follows. Paraconid height increases from M2-4 but is reduced in M.-,. The metaconid height in M 2.a is lower than in M4-3. Metaconid length at the base of the cusp decreases markedly from M_. 一 M :,. The The entoconids of M3 and M«arc subequal and larger than the entoconid on M2 which is larger than the entoconid on M5. The hypoconulids of are subequal in height. The protoconids of M2 and M5 are subequal and shorter than the protoconid of M3 which is subequal to the protoconid of M4. The hypoconid height decreases from lo M :,. The talonids of M2 一 Ms are narrower than the trigonids. The paracristid of M2 is shorter than that of M3 which is subequal to that crest in M4. The paracristid of is shorter than that of M,. The metacristid length increases from to M. The metacristid of Mr, is shorter than the metacristid of M4. The cristid

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9 VAN DYCK: ANTECHINUS PUTEUS, AN EXTINCT DASYURID 67 obliqua intersects the trigonid in a progressively more lingual position from M2 to Mr,. The hypocristids of M2 to arc subequal in length. The anterior cingulum length decreases slightly from M3 一 M 4. The posterior cingulum length decreases from Mjj _ M4 (absent in Mr>). The buccal cingulum length decreases from M2 M,. The length of the buccal cingulum of Mb is slightly greater than that of M4. The length of the greatly reduced lingual cingulum of M.i is greater than that of M4. DISCUSSION Antechinus puteus exhibits its closest affinities with A. flavipes. Because of its size, A. puteus is compared with the largest race of A. flavipes, A. /. rubeculus which ranges from Mt. Spec. (18 STS 146 ITE) to Home Rule (15 42 S #E) in northeastern Queensland. A. puteus displays a number of features which.. seem to be extraordinary developments of ordinary A. flavipes tendencies (Archer 1978: 72). Although molar measurements for F8445 (M2, Ma, M4) appear to stretch the limits of acceptable intraspecific variation, these arc statistically insignificant (see Tabic 1). However, several features of A. puteus arc unique; the single-rooted P3 which reduces the overall length of the premolar row; the extreme buccal orientation of the talonid endoloph from the base of the mctaconid to the hypoconulid; and the reduction of the paraconid of M2. Archer (1978) notes the outstandingly short nature of the premolar row in A. puteus. This is further demonstrated in values for Pra. and Ps in Tabic 1. More importantly, the relative length of the premolar row (compared to the length of the molar row) shows that A. puteus has a relatively much shorter premolar row than any other measured specimen of A. f. rubeculus ot A. f. flavipes. In A. puteus, the overall reduction of the prcmolar row is due to the effect of the obliquely oriented, single-rooted Ps. Archer (1976) notes that reduction of P3, and widening of the premolar tooth row, arc derived states within the dasyurids as a whole. Elsewhere (Van Dyck 1982) I have remarked on the derived condition of the crowded premolar row in A. flavipes as compared to the relatively uncrowdcd condition in A. stuartii. Associated with the compression of the premolar row in A. flavipes, there has been a posterior shifting of C 1 to accommodate a more massive Cj. Bcnsley (1903) related the tendency of simultaneous premolar reduction and canine enlargement to a need for increased killing efficiency in more carnivorous forms. Tate (1947) noted that short muzzles and short palates accompany short, crowded tooth-rows in vertebratekilling dasyurids, and Thomas (1887) related premolar reduction to specialization in dasyurids. A direct relationship in dasyurids between M-_. paraconid reduction and reduction in P3 has been noted by Archer (1976) who saw its effect as premolarizing the trigonid of Mj and hence shifting posteriorly the premolariform-molariform boundary. The reduction of P3 in A. puteus is accompanied by a reduction in the paraconid of M2to a low, broad shelf. The sharp buccal swing taken by the talonid endoloph (from the base of the mctaconid to the hypoconulid) of M2 M, in A. puteus is unparalleled by other antechinuses. Greater talonid reduction in A. flavipes has been used to distinguish it from A. stuartii (Van Dyck 1982), but in A. puteus a reduction much greater than that seen in A. flavipes occurs in the talonids of Ms and M. This is demonstrated in the comparison of Fig. 4b with Fig. 4d. Reduction of the talonid of Ma reflects a general dasyurid trend away from the ancestral condition as does also the reduction of the main talonid cusps from the ancestral three to one (Archer 1976). The more specialized nature of A. puteus is demonstrated in both the excessive reduction of talonid width and the reduction of Ms talonid cusps. It is likely that if maxillary remains of A. puteus are found, the protoconcs and paraconcs of the corresponding upper molars may be separated by a

10 68 AUSTRALIAN MAMMALOGY relatively shorter distance than that found in any other antechinuses. Antechinus puteus was probably con temporaneous with A. flavipes because fossil specimens o f both were recovered from the same deposit in Russcndcn Cave. These contem poraneous specim ens o f A. flavipes. however,are smaller than A. puteus (see Table 1 and Archer 1978), the difference being o f a com parable order as that between A. flavipes flavipes (southeastern Queensland) and A. flavipes rubeculus (northeastern Queensland) (see Van Dyck 1982). A C K N O W L E D G EM EN TS I am grateful to Dr. J. Erzetich, Brisbane, for supplying me with X-ray photographs of the paratype, F8445, and to Mr. R. Raven (Queensland Museum) for taking the electron micrographs. I would like to thank Dr. M. Archer (University of New South Wales) and Dr. R. W ells (Flinders University, Adelaide) for their critical reviews o f this paper. REFERENCES Archf.k, M., The dasyurid dentition and its relationships to that of diddphids, thylacinids, borhyacnids (Martupicamivora) and paramelids (Peramelina: Marsupialia). AuU. J. ZooL, Suppl_ Ser. 39: l-i4. A rch er. M., Quaternary vertebrate faunas from ihe Texas Caves of southeastern Queensland. Mem. Qd Mus. 19: A rch er. M., Review of the dasyurid (Marsupialia) fossil record, integration of data bearing on phylogenetic interpretation, and suprageneric classification. Pages in "Carnivorous marsupials", ed by M. Archer. Roy. Zool. Soc. New South Wales: Sydney. B e n sle y, B. A., On the evolution of the Australian Marsupialia; with remarks on the relationships of the marsupials in general. Trans. Linn. Soc. Lond. (Zool.) 9: &i-2v7. Ride, W. D. L., *'A guide to the native mammals of Australia." Oxford Univ. Press: Melbourne. Tate. G. H. H., Results of the Archbtild Expeditions. No. 56. On the anatomy and classification of the Dasyuridae (Marsupialia). Bull. Amer. Mus. Nat. Hiu. 88: Thom as On the homologies and succession of the teeth in Ihe Dasyuridae, with an attempt to trace the histor>' of mammalian teeth in general. Phil, irons. 128: Van D yck, S., The relationships of Antechinus uuartii and A. flavipes (Dasyuridae, Marsupialia) with special reference to Queensland. Pp in "Carnivorous marsupials", cd. by M. Archer. Roy. Zool. Soc. New South Wales: Sydney.