THE BRONZE QUOLL, DASYURUS SPARTACUS (MARSUPIALIA: DASYURIDAE), A NEW SPECIES FROM THE SAVANNAHS OF PAPUA NEW GUINEA
Stephen Van Dyck
Van Dyck, S.. 1987. The Bronze Quoll, Dasyurus spariacus (Marsupialia: Dasyuridac), a new species from the savannahs o f Papua New Guinea. Ausi. Mammal. II: 145-156.
$p. nov. is described from the low mixed savannahs of southwestern Papua New Guinea. It differs from all other species of D asyurus in the aircm e narrowness of the rostrum measured bcl^vcn left and right lachr>'mal canals. Its hallux and ear lengths are small. D. spartacus is a specialised species; its affinities lie with D. altwpunaaius and possibly D. du n m alli (a fossil species).
Key words: Dasyurus spartacus, marsupial, Papua New Guinea, phytogeny.
Stephen Van Dyck. Queensland Museum, PO Hox 300, South Brisbane, Queensland, Australia 4101. M anuscript received 16 July 1987.
UNTIL 1979 Dasyurus albopunctatus Schlegel, (MNHN). Additional specimens examined and was the only quoll known from Papua New Guinea. mentioned in this paper have registration numbers In that year J. Waithman published results o f a prefixed as follows: National Museum and Art mammal survey undertaken in the Trans-Fly Plains Gallery, Papua New Guinea, Boroko (PM); of southwest Papua New Guinea during 1972-3. Queensland Museum Brisbane (J, JM or F); Part of this collection included 5 specimens o f a Western Australia Museum, Perth (WAM); Dasyurus referred to D. g e o ffr o ii (Waithman 1979) Australian Museum, Sydney (M). collected at Morehead, Mibini and Mari. I have examined the following holotypes: Subsequent references to this Dasyurus have also Dasyurus geoffroii geoffroii Gould 1841 (BM been made under the specific title o f g e o ffro ii 41.1213), D. g.fortis Thomas 1906 (BM 6.8.1.340), (Ziegler 1977, Archer 1979, Archer 1982, Honacki, D. haUucatus hallucatus Gould 1842 (BM Kinman and Koeppl 1982, Arnold 1983, Taylor 42.5.26.16), D. h. exilis Thomas 1909 (BM 9.4.23.8), 1984). It is now dear that these specimens represent D. h. predator Thomas 1926 (BM 15.3.5.77) D. h. a new species of Dasyurus which is described here n esaeu s Thomas 1926 (BM 26.3.11.25), D. as D. spartacus. albopunctatus albopunctatus Schlegel 1880 (RMNH Specimen a.), D. a. fu scu s Milne-Edwards 1880 (MNHN 1880-1463), D. a. daemoneUus MATERIALS AND METHODS Thomas 1904 (BM 3.12.1.24), D. du n m alli Dasyurus systematics follow Kirsch and Calaby Bartholomai 1971 (F 6579). The holotypes o f D. (1977); terminology of cranial, external and dental viverrinus (Shaw 1800) and D. m aculatus (Kerr morphology follows Archer (1981); and tooth 1792) are presumably lost. number Tollows Archer (1978). Cranial and dental measurements were made with NSK electronic SYSTEMATICS digital calipers after the method shown in Table I. All specimens compared in diagnosis were adults Dasyurus spartacus sp. nov. (Fig. 1, Table 1) with fully erupted P ：. Holotype. PM 22000, adult male, dry skin, skull Specimens o f Dasyurus were examined in each and dentaries. collected 11 April 1973 by J. of (he following museums: Bernice Bishop Waithman. Museum, Honolulu (BBM); British Museum Type locality. Morehead, Trans-Fly Plains, Papua (Natural History} London (BM); Rijksmuseum van New Guinea 8041*S, 141。39^ (Fig. 2>. Natuurlijke Histoire, Leiden (RMNH); American Museum of Natural History, N w York (AMNH); Paratypes. From Morehead, adult male PM22004 Museum National DTIistoire Naturelle, Paris in ethanol with skull extracted, collected 25 146 AUSTRALIAN MAMMALOGY
November 1973 by H. Parnaby; juvenile male PM22001, puppet skin with skull extracted, collected 17 May 1973 by J. Waithman. From Mari (oil line Na 1) 90U S, 141°42 E, adult female PM22003 skull only, collected 18 July 1973 by J. Tameiona. From Mibini 8°50*S, 14l°38rE, juvenile female PM22002 skin with skull extracted, collected 26 June 1973 by J. Waithman. Diagnosis. D. spartacus is a medium sized but thick set Dasyurus immediately separable from all other species of D asyuru s by the extreme narrowness of the rostrum measured between left and right lachrymal canals (Fig. 3). It also possesses a reduced hallux (Fig. 4) and the pinna is small. Externally. Unlike D. maculalus, D. spartacus lacks white spots on the tail and lacks striate pads on the fore and hind feet. D. spartacus differs from D. maculatus in being much smaller (eg. head-body length adult male D. maculatus (J8059, in absence | | o f holotype) 605 mm, D. spartacus (holotype s l PM22000) 345 mm, and having much smaller white spots on the body (average diameter o f side spots in D. m aculatus (J8059) 34.4 mm, in D. spartacus 2 I (holotype PM22000) 5.4 mm. i i D. 5/w/7“c*u5 differs from Ww/vVjiw (Shaw, 1800) in having a hallux. D. spartacus differs from D. geof/roii in having 2 a body colour a deep bronze to tan brown with a | a black tail and much smaller white body spots ^ (average diameter o f side spots in D. geof/roii 1" (holotype BM41.1213) 12.1 mm, in D. spartacus (holotype PM22000) 5.4 mm. D. spartacus possesses a vestigial hallux (Fig. 4) which is approximately y〇 5 58% smaller than that in D. g e o ffr o ii (eg. D. spartacus (spirit specimen PM22004) 2.1 mm, D. r g e o ffr o ii (spirit specimen JM2057) 4.9 mm. D. spartacus has much smaller ears (eg. D. spartacus 2.1 (holotype PM22000) 30.0 mm, D. g e o ffr o ii II (holotype BM41.1213) 60.0 mm. D. spartacus lacks the striate pads on fore and hind feet present in D. hallucatus and D. IIo 多 albopunctatus. D. spartacus also differs from D. hallucatus and D. albopunctatus in being larger (eg. II head-body length adult male D. hallucatus (JM 5701, in absence o f measurements accompanying i s holotype) 285 mm, D. albopunctatus (M14853, in absence of measurements accompanying holotype) 266 mm, D. spartacus (holotype PM22000) 380 mm. Internally. D. spartacus differs from D. maculatus in being much smaller (adult male basicranial length D. maculatus (J8935, in absence of holotype) 96.7 VAN DYCK: A NEW DASYURUS 147
Fig. /. Dasyurus spariacus, skull and dentary o f holotype PM22000; scale, a x 0.80, b x 0.80, c x 0.83, d x 1.12.
D. spartacus differs from D. hallucatus in being much larger (basicranial length adult male D. hallucatus (J16752 as holotype is female) 62.5 mm, D. spartacus (holotype PM22000) 70.2 mm, having a *U* shaped incisor row, having the metacone of M2 posterior to stylar cusp D, having M5 reduced with no metacone, having metacrista of M3 shorter than the metacrista of M4, having metaconids reduced and having a short and crushed premolar row with bulbous premolars. D. spartacus differs from D. albopunctatus in being larger (adult male basicranial length D. albopunctatus (M14854, as holotype is female with smashed skull) 57.0 mm, D. spartacus (holotype PM22000 ) 70.2 mm, and in having less reduced Fig. 2. Map showing collection sites (Morchead. Mibini metaconids. and Mari} of Dasyurus in Papua New Guinea. Description. A thick-set, narrow-nosed quoll mm, D. spartacus (holotype PM22000) 70.2 mm. distinctive for its rich dark, golden-brown ionings, D. spartacus also has a procumbent and separated minute white spots, dark golden-bronze feet and I1, a metaconid on M2 and a short, crushed dark tan tail. premolar row. Pelage. Colour for holotype as follows (colour D. spartacus differs from D. viverrinus in having names after Ridgway 1912); dorsal fur above a procumben( and separated I1, greatly reduced shoulders 11.2 mm with basal 4.5 mm fine, silky, metaconids, and a short and crushed prcmolar row. apical 6.7 mm medially thickened, spinous. Basal 8 mm Clove Brown (dark chocolate), median 1.6 D. spartacus differs from D. geoffroii in having mm Ochraceous-Tawny (orange-brown), apical 1.6 a short and crushed premolar row with broad, mm Fuscous Black. Fur of back between shoulders bulbous premolars. appears as dark chocolate flashed with orange. 148 AUSTRALIAN MAMMALOGY
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Clove Brown extends anteriorly to tip o f nose and eye so that sides o f face contrastingly light against posteriorly to tip o f tail. On head, relatively long dark head-patch. hair (10 mm) lacks lawny band thereby creating very dark (Clove Brown) triangular head patch from H air on rum p (to 9 m m long) sim ilar co lo u r to nose, above eyes, back to between ears. Hair shoulders but median Ochraceous-Tawny band betw een eye and car and for 4 m m above each eye slightly longer (2.5 mm). Sides o f body, forearms, has median band (up to 20 mm long) Chamois thighs characterised by increase in length o f median (light yellow). Band may increase to 4 mm below colour band (Yellow Ochre), to 6 m m (on hair o f VAN DYCK: A NEW DASYURUS 149
II mm) so overall colour of sides is bright golden- Tail. Tail shorter (by approx. 20%) than nosc-vent bronze. Belly fur soft, to 21 mm between hindlegs, length, non-incrassatcd, bushy toward tip. Cream Colour (almost colourless). H indfoot. Interdigital pads raised, joined, "Hiil short-haired for basal '/j, more distally hair minutely granulated. Apical granules only slightly lengthens to 37 mm at tip and darkens to Fuscous larger than surrounding granules. Slightly enlarged Black (black) at tip. Tail shorter haired and lighter granule may represent hallucal granule on both feet. coloured ventrally. Hair on upper surface hind and Metatarsal granules not present in holotype. Hair forefoot, a shiny Sepia. on foot covers heel. Terminal pads of digits weakly striate (Fig. 4). Small white spots (maximum diameter 6.5 mm) occur randomly from forehead to rump. No white Ear. Pinna very small, curled external edge on spots on ventral surface or on tail, fore or hindfeet. large supratagus. Fawn hairs on postcro-intcrnal and ventral margins o f pinna. Vibrissae. Approx. 25 mystacial vibrissac (up to 35 mm) each side in 5 distinct horizontal rows. More D entition (Fig. 1). Upper incisors: I1 narrow, dorsal vibrissae Fuscous Black, more ventral peg-like, procumbent, slightly curved, taller- colourless; supra-orbital vibrissae (Fuscous Black) crowned than all upper incisors; 11 separated by number 2 left, 3 right; genals (Fuscous Black and large diastema from I2. Left and right 11 widely colourless) 6 left, 5 right; u!na-carpals (colourless) separated. Crown height, length subequal in I2 and 3 right, 6 left; submentals (colourless) 6. I3.14 slightly greater crown height and length than 150 AUSTRALIAN MAMMALOGY
I2 and I3. All upper incisors lack buccal cingula yet cingulation. Crown height P2 much taller P 卜 no lack o f differentiation between root and crown. Prcmolars relatively broad, bulbous, both with I4 with no anterior or posterior cusp. Root of I4 weak posterior cusp, P2 with very weak anterior narrow. cusp. Bulk of each premolar mass concentrated posterior to line drawn transversely through middle Upper canines: C 1 long, slender, caniniform of ihe two premolar roots. with indistinct boundary between root and crown. No buccal or lingual cingula. No anterior or Lower molars: All molars broad, bulbous, tightly posterior cusp. crushed together. M2 talonid wider than trigonid, anterior cingulum absent. No buccal or lingual Upper prcmolars: Small diastema between cingulum. Paraconid absent. Mctaconid reduced to crowns C* and P*, P 1 and P2, P2 and M2. P 1 small bulge o f enamel subequai height 10 minute shorter crown height than P2. Minute anterior, entoconid. Metacristid greatly reduced, oblique to posterior cingula cusps P 1 and P2. P2 broad long axis of dentary; hypocrislid more postero-lingually. perpendicular. Cristid obliqua very short, extends Upper molars: Posterior tip P' not contacting from hypoconid to posterior wall of (rigonid parastylar corner M2. Anterior cingulum below intersecting at point well lingual to point directly combined stylar cusp B-paraconc short, broad, below tip of protoconid. Hypocristid terminates complete to prominent paraconulc at base of midway between hypoconid, metastylid. Long, low paraconc-stylar cusp B apex. Paracone-stylar cusp entoconid. From base of meiaconid posteriorly B o f M 2 approx, half height o f mctaconc. Stylar entoconid forms prominent semicircular bulge of cusp C, E not visible on L or RM2. Trigon basin enamel (occlusal view) on cndoloph. Indistinct plunges steeply from prominent metaconule down posterior cingulum. to weak posterior cingulum. M 3, talonid slightly wider than (rigonid. M3, stylar cusp B and paraconc separate. Broad Anterior cingulum very poorly developed originating lingually in weak parastylid notch into anterior cingulum contacts metastylar corner M2, tapers down along base of paracrista, rises steeply which hypoconulid tucked. No buccal to base of paraconc apex at prominent paraconule. cingulum. Narrow, weak posterior cingulum almost M3 lacks stylar cusps A, C and E. Stylar cusp D obliterated by crushing anterior cingulum o f M4. prominent and broad. No posterior cingulum; Paraconid well developed, smallest trigonid cusp. prominent metaconule as in M2. Entoconid well developed, subequal in height to paraconid. Cristid obliqua extends from M4, anterior cineulum complete, broader, much hypoconulid to posterior wall of trigonid longer than M ' Paraconule, metaconule intersecting trigonid at point directly below tip of prominent. Stylar cusp D reduced but broad. Stylar protoconid but well buccal 10 metacristid fissure. cusps C, E absent. Posterior cingulum absent. From base metaconid posteriorly, endoloph follows line oblique and lingual to dentary axis. M5, broad at anterior cingulum terminates quickly away from metastylar corner M4, posterior M4, trigonid wider than talonid. Prominent cingulum absent. Protocone reduced, narrow. parastylid wraps around hypoconulid M3, weak Metastylar corner reduced. anterior cingulum on M4. No buccal or posterior cingula. Reduced cristid obliqua intersects trigonid Lower incisors: I t much taller crown height than at point well lingual to longitudinal vertical midlinc h- *i* *2» b 〇val *n anterolateral view, I( and I2 drawn through tip 〇r protoconid but buccal to gouge-like in occlusal. I2 subequal crown height I3. metacristid fissure. Entoconid M4 worn but was All lower incisors with prominent lingual cusp. I3 obviously large, making prominent bulge of enamel with most conspicuous posterolingual cusp almost on endolph. Rest o f m orphology as in M j except as (all as primary incisive cusp. Lower canine rests that metaconid is more reduced. between posterior cusp and primary incisive cusp *3- M5, (rigonid much wider than talonid. Anterior Lower canines: Q large, broad, caniniform, cingulum as in Posterior cingulum absent. Of characterised by upward, forward projection and three main trigonid cusps metaconid slightly taller than paraconid but both dwarfed by protoconid. sickle-like curvature root to crown tip. No buccal, Hypoconid M 5 talonid much smaller than \14. very weak lingual cingulation, no posterior cusp. Between hypoconid and base of metacristid, cristid Lower premolars: Premolar row very short, Pj, obliqua forms low, very weak cresi, which contacts P2 crushed together, overlapping, separate from trigonid wall directly below meiacrislid fissure. M2 by small diastema. Lack buccal, lingual Evidence o f small entoconid. VAN DYCK: A NEW DASYURUS ISI
S k u ll (Fig. 1). D. sp a r ta c u s is unique amongst holotype or paratypes. The tail is short-furred, but D a sy u r u s species in having a high skull and demonstrates the golden base which is replaced by extremely narrow rostrum between the lachrymal black fur one third o f the way down the tail toward canals. Sagittal and nuchal crests are highly the tip. In spirit paratype adult m ale PM22004 developed. Dorsally the rostrum is gently grooved Fuscous black hairs appear slightly bleached to dark longitudinally by a depression running.along the reddish brown. The partly e\crtcd penis shows the nasal sutures. One lachrymal foramen occurs on penis appendage observed and mentioned by each side of the orbital rim. The left and right Woolley and Webb (1977). alisphenoid tympanic bullae are widely separated Reproduction. Waithman (ncld notes) recorded and relatively small. The foramen pseudovale is small and bisected by a (hin bridge of the 7 nipples for PM22003. A ssum ing growth rates alisphenoid. The foramina o f the transverse canal similar to those recorded for Dasyurus viverrinus are large and obvious, whereas the cntocarotid (Fleay 1935), juveniles PM2200I and PM22002, collected 17 May 1972 and 26 June 1973 respectively, foramina are small and obscure. The internal jugular canal foramina are large; the canals are may have both been born in February. raised and prominent. The posterior lacerate Habitat. Specimens of Z2 were collected foramina are large and exposed as are the carotid in low mixed savannah at three localities: three from foramina. The premaxillary vacuities extend from Morehead 8041’S* 141039*E (Rg. 6>; one from Mibini the level of the I4 root back to the level of the 8°50'S, I41°38E; and on e from Mari 9°irS, middle of the C 1 root. The very small maxillary 141°4rE. PM22001 was "brought in by sch ool vacuities extend from (he level o f the protoconc root children from a road pit*(field notes J. Waithman) o f M2 back to a level slightly posterior to the and PM10400 was killed while raiding a fowl house. protoconc root of M4. Minute palatine vacuities Colleccion details for the other specimens are not extend from the level of stylar cusp B on M5. available. Collection areas experience a monsoonal Variation in paratypes. Dental and cranial. In climate where 75% o f annual rainfall (1682 mm for adult female PM22003 palatine vacuities are large Morehead) falls during a wet season lasting from and more numerous than in the holotype. All (eeth Decem ber 10 May (Paijmans, Blake, Blecker and M cAlpine 1971). are very worn. The anterior cingulation of M5 is broad and complete 10 the trigon basin. A very wide The habitat type, low-mixed savannah (Paijmans diastema separates Pj and M2. In juvenile female et al. Plate 16, Fig. 2, 1971) occurs in Indorodo, G o e PM22002 R and l.M4"are partially erupied, R and and Mibini land systems where the habitat is LM 5 are unformed. RM 2 shows large slylar cusp inundated and waterlogged during the wet season C. LM> shows bifid enloconid while L and RM 5 and burned during the dry season. Trees average are pariially erupted. The unworn dentition o f this approximately IS m in height but som e grow as tall paraiypc clearly demonstrates the very reduced as 28 m. Most common species include T ristan ia metaconids of R and LM 2 and the strong suaveolans, Melaleuca symphocarpa, development of entoconids on M 2_5. Xanthostemon crenulalus, Grevillia glauca and In juvenile m ale PM22001 M445 are unformed. Banksia denlata. Less common species include All other teeth arc partially erupted. Stylar cusp C Deplanchea leiraphylla, M. viridiflora, E. is present on both R and LM^ but not M3. The crown tip of RP2 is bifid and a minute, single- rooted, splinter-like M 1 occurs between RP2 and RM2. On the dentary all teeth arc partially erupted, M 5 is unformed. The entoconids o f R and LM 2 are bifid.
In adult male PM22004 all teeth are so worn that RP2 is represented by two blunt roots, the tooth crown (presumably) having worn away.
External characters. In juvenile paratype female PM22002 (Fig. 5) both hind feet and the left forearm are missing. Colour closely matches holotype cxcepi that black guard hairs appear much more prominently ovxrr the back and sides. Juvenile paratype male PM2200I is very young and newly flfg.又 Juvenile female (PM22002>. furred and appears more golden in colour than the Phoio: J. Waithman. 152 AUSTRALIAN MAMMALOGY
premolars generously spaced. In N w Guinea ihesc premolars arc crowded and crushed together in a short row.
Uncrowded premolars have long been considered (he primitive condition in didelphoids and hence Dasyuroids (Thomas 1887, Bensley 1903, Tale 1947, Archer 1976). But although premolars in the genus D a sy u ru s are generally widely spaced, the loss o f P3 indicates considerable prelusive specialisation. It could well be that (he relaxed condition o f the two-premolared row is a further derivation in the dasyurid pattern o f premolar crushing and single- rooting o f P3 prior to its ultimate expulsion from the row. This pattern is demonstrated in Table 2, where in the more primitive condition eg. M u rex ia /owgfcflMc/a/ff, th elow er prem olar row may occu py up to 32% o f the entire lower tooth row. In Phascogale tapoata/a, a slightly more specialised dasyurid exhibiting a trend toward loss o f P 3, the value decreases to approximately 25%. In Sarcophilus harrisii where P 3 has been lost from both upper and lower tooth rows, the percentage Fig. 6. Dasyurus spartacus habitat, low-mixed savannah a( Morchcad, Papua New Guinea. Pholo: H. Parnaby. o f Icmw premolar r w drops to approximately 17%. In D. hallucatus, D. viverrinus, D. m aculaius and polycarpa, T. longivalvis. Acacia spp., D ille n ia D. g e o f f r o ii this value ranges from approximately alala, Parinari nonda and Xanthosiemon barassii. 22-23% and suggests a trend o f prcmolar loss then A patchy open shrub layer 1.0-3.3 m in height subsequent rostrum and dentary lengthening — a contains Acacia leptocarpa, Choriceras tricorne, rc\'ersal o f the trend exhibited by Sarcophilus. The M etasioma polyanthum, Rhodamnia, Acacia simsii, greatly reduced premolar rows o f D. spartacus and Timonius dlochidion, Sinoga tysicephala and \ow D. a lb o p u n c ia tu s (19-20??) appear to dem onstrate P an dan us. The ground layer (with a cover of the prcmolar condition just subsequent to the loss 80-100%) contains Imperaia cylindrica, o f P3 and would suggest that these two species were Pseedopogonotherum irriians, Germainia capilata, in Tact more primitive than all the others. Eriachre squarrosa, Schoenus spp., S c h le r ia and The possibly Pliocene D a sy u ru s d u n m a lli from Rhynchospora rubra. Nepenthes and D a n e lla are the Chinchilla Sand, with its single rooted P3 could always present and Ulricularia, Drosera and lend support to this argument. With a crushed and E r io c a u io n occur in areas o f very poor drainage only slightly greater proportion o f premolar row (Paijmans et al. 1971). (22%), D. dunmalli could represent the species Elym ol〇K>'. The name sp a r ta c u s embodies many ancestral to D. spartacus and D. albopunciatus. features shared by this dasyurid with the notorious However, (he sam e constraints that Bartholom ai Thracian gladiator — strength, a tenacious fighting (1971) and Archer (1982) applied to the spirit, and the capacity to draw blood. interpretation o f the significance o f D. dunmalli in D a s y u r u s phylogeny must be applied DISCUSSION simultaneously to D. spartacus an d D. albopunciatus. In these two species a consideration Past reference to (his new dasyurid as D. g e o f f r o ii o f character states apart from the prcmolar row has resulted from not only its strong supcrHcial indicates specialisation of a highly derived nature resemblance lo D. geoffroii but also from a eg. disproportionate C 1, V-shapcd upper incisor reasonable expectation that D. geoffroii with its row, m etacone M 2 perpendicular to stylar cusp D, extensive pre-European distribution throughout M 2 reduced without entoconid, specialised penis, Australia (Ride 1968, Archer 1979, Johnson and metacrista M 3 shorter than in M4, greater R off 1982) could occur in New Guinea. One notable reduction o f paracones, metaconids reduced and feature, however, differentiates quolls north and upper premolars bulbous (see Tabic 3). south o f Torres Strait; the condition o f the premolar row. In Australia the genus D a s y u r u s is These fea⑴res clearly make D. spar/acus and Z). characterised by a long premolar row with albopunciatus (along with D. dunmalli for fw e r VAN DYCK: A NEW DASYURUS 153 reasons) inclligiblc for consideration as the most z r s 13 s.f £ I S S s 9 s 69 p plesiomorphic members o f the genus when the > 卜 oo . u . 对 5. r £ morphologically primitive D. h allu ca tu s is »n r- 08.0 l r 6 卜 considered. If D. dunmalli was considered the sister l l£ s 0 3Z$ z s s 6 5 species of D. spartacus and D. albopunctatus, then o 0 s o 00• 0 JGIOEoJd I -R.9 IZ T 8 06TZ 5.0Z-S.61 loss of P3 must have occurred once in D. hallucatus SS 9 69s-9r91 8 S 卜 € . r and then reversal o f the premolar character state . S S c 001 § - c - - r cx:curred once in the D. dunmalli lineage whereupon - P «s- 9 8 -s 穿 00T the persistent Dasyurus 2-premolar trend reemerged 6 . S J 00 00 - . 03 6 in D. albopunctaius and D. spartacus (Fig. 7A). o Z i 1 ZZ s S i l I n o r o s o o ^ o r c While this is a tempting and parsimonious approach 70 S z S z 02K S 2 1 o . 9 2 . ^ OOZZ it assumes many synapomorphies from D. dunmalli A O0 0 P USZC 如 广 o f which so little material (a few incomplete M-M o 2 0 91Z -M eop SS6 r l - dentaries) is available. i l . oo.9l 6 s 1 Archer (1982) has suggested that loss of P3 may 09f £r>8 S 1 8 I 3 2 have occurred twice within the genus, once from A
1 6 C - Z 0 6 -9 1 - .6 2 0 . 6 - 3 U o f the species vi’ve/r//i叫 容 and moew/a/ui：. 8 z 9 . s _ le r c. 9QO 2 S . . S 卜 .2 However, two synapomorphies (more reduced J § - 62 S - I 7 9. 卜 C metaconids and broad, oval premolars) of dunmalli l i Z o . SII M 卜 e S S 2 o 1 and the spartacus-albopunctatus-maculatus group l 001 q .2爸 suggest that D. dunmalli is more closely related to ? 9 - a 0 9 62 ^ 0 R 62 l Oo £ 0 8 0€ 9 8 l l s these three species than to viverrinus and geof/roii. oo 0 o d KU!se3J3u! uv H o o M ■»0 卜 二 _ M Mo i In this case loss o f P3 may have occurred once in l M 5 C6>C 9 19.99 -Ms 6 9 -H2 £ the hallucatus-viverrinus-geof/roii line and then -H 5. . l r -m9 0. | o 2 u S O0 lx 2 once again in (he ancestor (D. dunmalli?) o f D. z S 9 Z spartacus and D. albopunctaius. p; C 6 6 > 6 8 32 ll.ll 6 6 01 JCl!M A . s Soo . If consideration is given to Bartholomai's (1971) u d oo1 C2 r»080 warning regarding the dubious nature of speculating 6 a: s 的 l s ) . s 05 s n e(upnp;>J 1 K - on the relationships o f D. dunmalli then another 80 0o o .o s . 0 o o approach (free from the influence of D. dunmalli) J is represented in Fig. 7B. To opt for the most r 6 9 . 9 - 2 moj JSOUKUd s c. - z r 2 「 5 8 - parsimonious approach the assumption in this case § "If 卜 - s . CJ- T 2 - 6 oc- r v? J is made (hat the crushing o f the premolar row in “ 8 -> R oo. r I I D. spartacus and D. albopunctatus is a highly 0J 00 oo r* O M 'c l derived condition associated with the shortening of 0 ou the rostrum. Archer (1976) notes that the reduction I J d JC S o f P3 and widening of the premolar tooth row are 1 I -oo OOU J n derived states within the dasyurids as a whole. 7 A Bensley (1903) related the tendency of simultaneous 1 S rtp premolar reduction and canine enlargement to a jo need for increased killing efficiency in more carnivorous forms. Tate (1947) noted that short l l l muzzles and short palates accompany short, scu crowded tooth rows in vertebrate-killing dasyurids II qsuB s «i3 and Thomas (1887) related premolar reduction to s - > o n l ss o specialisation in dasyurids. In this interpretation, p s 3 s iid z M m D f} sn u fllo n / a J z i3 f i ff£ J O a afuuopoDUJ :s u n u D. spartacus and D. albopunctatus are sister species u uD u n ^ f i a l s d a jo more closely related to one another than either is o a .S o a 'sUJXq a JJ :s D l q aJ J a U5UGJ to D. geof/roii. l I f E 鉍 - > > d d The upshot o f these considerations is twofold. s Je. s Firstly that regardless of a preferred phylogeny, D. . *n 4«! js 154 AUSTRALIAN MAMMALOGY Fig. 7. Two cladograms of the hypoihetical relationships of Dasyurus spartacus. Cladogram A includes the Pliocene species D. dunmalli and reflects interpretaiion 4A of Tabic 3. Cladogram B includes only extant members of ihe genus and rcflccis imcrpreiafion 4B o f Table 3. See Tabic 3 for reference to all other apom orphies depicted in the cladograms. VAN DYCK: A NEW DASYURUS 155 1. l ' narrow, peglike P P A P P P - 2 . I- procumbent, separate P P A P P A - 3 . cl proportionate P A A A A k A A3 A3 A3 A? A2 A. Premolar row 1ongv prem oUrs Spdced ^ i n TT TT TT TT 5 . Prem olars narrow, sm^ll p P P A A A A 6 . M olars non-bulbous p P P P P A P 7 . In cisor row V-shaped p A A A A A - 8 . Paricooes reduced A1 A2 A2 A? A2 A? - 9 . M etaeonids unreduced P A2 A1 A? A3 A3 A2 1 0 . Hetaconc H? perp. to St. cusp D P A A A A A - 1 1 . Hetacrista equal to or longer P A A A A A - 1 2 . H- unreduced P A A A A A - 1 3 . w ith metacone P A A A A A - 1 4 . with eDtoconld A A A A A P - 1 5 . Skull height low P P P A P P - 16. Broad rostrum width between lachrym als P P P A P P - 1 7 . Peon sim ple P A A A A A - 1 8 . Possession of hallux P P A2 A1 P P - 1 9 . Hind foot short and broad P P A P P P • Table 3. Character states of features thought to be useful in phylogenetic interpretation of quoll evolution. Character 4 presents two interpretations (sec discussion), a. (hat the premolar slate in D. dunmalli represents the least derived of all conditions and b. that excluding D. dunmallL the crushed premolar stace in D. spanacus and D. albopunctaius represents the most derived premolar condition seen in the genus. sp a rta cu s is a highly specialised species, more ACKNOWLEDGEMENTS derived than D. g e o f f r o ii but not as specialised as D. maculalus. Its greatest affinities are with D. The examination o f type material held in overseas albopunctaius. Secondly that D. albopu nctaiu s, far institutions was made possible through the financial from being the traditionally accepted primitive support of the Royal Zoological Society of New species so frequently associated with D. h a llu ca lu s South Wales, the Linnean Society o f New South is one o f the more highly derived members o f the Wales and the Queensland Museum Board of genus. Trustees. I gratefully acknowledge the help o f the following for allowing me to examine specimens in One could have hoped for a more com fortable their care: J. Menzics (National Museum and Art phylogenetic scenario from the tropical rainforests Gallery, Papua New Guinea), L. Gibson (Australian of New Guinea out of which such a parade of Museum), D. Kitchener (Western Australian primitive laxa has emerged in the past. Museum), C. Kemper (South Australian Museum), 156 AUSTRALIAN MAMMALOGY A. Allison and C. Kishinami (Bernice Bishop marsupials in general. Trans. Linn. Soc. Lond. (Zool.). Museum, Hawaii), M. Tranier (Museum National 9:83-217. D'Histoire Naturelle, Paris), C. Smccnk Fleay, D., 1935. Breeding o f Dasyurus viverrinus and (Rijksmuseum van Natuurlijkc Histoire, Leiden), general observations on the species. J. M amm. 16(1): P. Jenkins (British Museum, Natural History, 10-6 . London), R. Angcrmann (Museum fur Naturkunde Honacki, J. H., Kinman, K. E. a n dKoeppl, J. W., 1982. dc Humboldt-Universititat zu Berlin) and G. Musscr Mammal species o f the world. Allen Press Inc. and (American Museum of Natural History, New York). The Association of Sysiematics Cotlections: 1 also thank H. Parnaby (University o f New South Lawrence, Kansas, USA. Wales) and J. Waithman (Conservation Johnson, K. A. and Roff, A. A., 1982. The Western Commission o f The Northern Territory) for Quoll Dasyurus geof/roii (Dasyuridae, Marsupialia) collection details and photos o f specimens, and in (he Northern Territory: hi&iorical records from particularly M. Archer (University o f New South venerable sources. Pp. 221-226 in Carnivorous m a rsu pials cd by M. Archer. Roy. Zool. S o c New Wales) and R. Raven (Queensland Museum) for South Wales: Sydney. their helpful discussions with me. B. Cowell (Queensland Museum) produced the photographs. Kirsch, J. A. W. and C ai.aby, J. H., 1977. The species 〇r living marsupials: an annotated list. 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