THE BRONZE QUOLL, DASYURUS SPARTACUS (MARSUPIALIA: …bionames.org/bionames-archive/issn/0310-0049/11/145.pdf · spartacus (spirit specimen PM22004) 2.1 mm, D. geoffroii (spirit specimen

TH E BRONZE QUOLL, DASYURUS SPARTACUS (MARSUPIALIA : DASYURIDAE), A NEW SPEC IE S FROM TH E SAVANNAHS OF

PAPUA NEW GU INEA

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 o f southwestern Papua New Guinea. It differs from all other species o f Dasyurus in the aircm e narrowness o f 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. dunmalli (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. Manuscript received 16 July 1987.

UNTIL 1979 Dasyurus albopunctatus Schlegel, was the only quoll known from Papua New Guinea. In that year J. Waithman published results o f a mammal survey undertaken in the Trans-Fly Plains o f southwest Papua New Guinea during 1972-3. Part o f this collection included 5 specimens o f a Dasyurus referred to D. geoffroii (Waithman 1979) collected at Morehead, Mibini and Mari. Subsequent references to this Dasyurus have also been made under the specific title o f geoffroii (Ziegler 1977, Archer 1979, Archer 1982, Honacki, Kinman and Koeppl 1982, Arnold 1983, Taylor 1984). It is now dear that these specimens represent a new species o f Dasyurus which is described here as D. spartacus.

MATERIALS AND METHODSDasyurus systematics follow Kirsch and Calaby

(1977); terminology o f cranial, external and dental morphology follows Archer (1981); and tooth number Tollows Archer (1978). Cranial and dental measurements were made with NSK electronic digital calipers after the method shown in Table I. All specimens compared in diagnosis were adults with fully erupted P：.

Specimens o f Dasyurus were examined in each o f (he following museums: Bernice Bishop Museum, Honolulu (BBM); British Museum (Natural History} London (BM); Rijksmuseum van Natuurlijke Histoire, Leiden (RMNH); American Museum of Natural History, N w York (AMNH); Museum National DTIistoire Naturelle, Paris

(MNHN). Additional specimens examined and mentioned in this paper have registration numbers prefixed as follows: National Museum and Art Gallery, Papua New Guinea, Boroko (PM); Queensland Museum Brisbane (J, JM or F); Western Australia Museum, Perth (WAM); Australian Museum, Sydney (M).

I have examined the following holotypes: Dasyurus geoffroii geoffroii Gould 1841 (BM 41.1213), D. g.fortis Thomas 1906 (BM 6.8.1.340), D. haUucatus hallucatus Gould 1842 (BM 42.5.26.16), D. h. exilis Thomas 1909 (BM 9.4.23.8), D. h. predator Thomas 1926 (BM 15.3.5.77) D. h. nesaeus Thomas 1926 (BM 26.3.11.25), D. albopunctatus albopunctatus Schlegel 1880 (RMNH Specimen a.), D. a. fuscus Milne-Edwards 1880 (MNHN 1880-1463), D. a. daemoneUus Thomas 1904 (BM 3.12.1.24), D. dunmalli Bartholomai 1971 (F 6579). The holotypes o f D. viverrinus (Shaw 1800) and D. maculatus (Kerr 1792) are presumably lost.

SYSTEMATICSDasyurus spartacus sp. nov. (Fig. 1, Table 1) Holotype. PM 22000, adult male, dry skin, skull

and dentaries. collected 11 April 1973 by J. Waithman.

Type locality. Morehead, Trans-Fly Plains, Papua New Guinea 8041*S, 141。39^ (Fig. 2>.

Paratypes. From Morehead, adult male PM22004 in ethanol with skull extracted, collected 25

146 AUSTRALIAN MAMMALOGY

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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 o f Dasyurus by the extreme narrowness o f 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 PM22000) 345 mm, and having much smaller white spots on the body (average diameter o f side spots in D. maculatus (J8059) 34.4 mm, in D. spartacus (holotype PM22000) 5.4 mm.

5/w/7“c*u5 differs from Ww/vVjiw (Shaw, ) in having a hallux.

D.1800

2D. spartacus differs from D. geof/roii in having

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/roiiy1" (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 58% smaller than that in D. geoffroii (eg. D. spartacus (spirit specimen PM22004) 2.1 mm, D. geoffroii (spirit specimen JM2057) 4.9 mm. D. spartacus has much smaller ears (eg. D. spartacus (holotype PM22000) 30.0 mm, D. geoffroii (holotype BM41.1213) 60.0 mm.

D. spartacus lacks the striate pads on fore and hind feet present in D. hallucatus and D. albopunctatus. D. spartacus also differs from D. hallucatus and D. albopunctatus in being larger (eg. head-body length adult male D. hallucatus (JM 5701, in absence o f measurements accompanying holotype) 285 mm, D. albopunctatus (M14853, in absence o f 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 o f holotype) 96.7

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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.

Fig. 2. Map showing collection sites (Morchead. Mibini and Mari} o f Dasyurus in Papua New Guinea.

mm, D. spartacus (holotype PM22000) 70.2 mm. D. spartacus also has a procumbent and separated I1, a metaconid on M2 and a short, crushed premolar row.

D. spartacus differs from D. viverrinus in having a procumben( and separated I1, greatly reduced metaconids, and a short and crushed prcmolar row.

D. spartacus differs from D. geoffroii in having a short and crushed premolar row with broad, bulbous premolars.

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 o f 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 metaconids.

Description. A thick-set, narrow-nosed quoll distinctive for its rich dark, golden-brown ionings, minute white spots, dark golden-bronze feet and dark tan tail.

Pelage. Colour for holotype as follows (colour names after Ridgway 1912); dorsal fur above shoulders 11.2 mm with basal 4.5 mm fine, silky, apical 6.7 mm medially thickened, spinous. Basal 8 mm Clove Brown (dark chocolate), median 1.6 mm Ochraceous-Tawny (orange-brown), apical 1.6 mm Fuscous Black. Fur o f back between shoulders appears as dark chocolate flashed with orange.


C love Brown extends anteriorly to tip o f n ose and posteriorly to tip o f tail. O n head, relatively lon g hair (10 mm) lacks lawny band thereby creating very dark (Clove Brown) triangular head patch from nose, above eyes, back to between ears. Hair between eye and car and for 4 mm above each eye has m edian band (up to 20 mm long) C h am ois (light yellow). Band may increase to 4 mm below

eye s o that sides o f face con trastingly light against dark head-patch.

Hair on rum p (to 9 mm long) sim ilar co lou r to shou lders but m edian Ochraceous-Tawny band slightly lon ger (2.5 mm). S ides o f body, forearms, thighs characterised by increase in length o f median co lou r band (Yellow Ochre), to 6 mm (on hair o f

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II mm) so overall colour o f sides is bright golden- bronze. Belly fur soft, to 21 mm between hindlegs, Cream Colour (almost colourless).

"Hiil short-haired for basal '/j, more distally hair lengthens to 37 mm at tip and darkens to Fuscous Black (black) at tip. Tail shorter haired and lighter coloured ventrally. Hair on upper surface hind and forefoot, a shiny Sepia.

Small white spots (maximum diameter 6.5 mm) occur randomly from forehead to rump. No white spots on ventral surface or on tail, fore or hindfeet.

Vibrissae. Approx. 25 mystacial vibrissac (up to 35 mm) each side in 5 distinct horizontal rows. More dorsal vibrissae Fuscous Black, more ventral colourless; supra-orbital vibrissae (Fuscous Black) number 2 left, 3 right; genals (Fuscous Black and colourless) 6 left, 5 right; u!na-carpals (colourless) 3 right, 6 left; submentals (colourless) 6.

Tail. Tail shorter (by approx. 20%) than nosc-vent length, non-incrassatcd, bushy toward tip.

Hindfoot. Interdigital pads raised, joined, minutely granulated. Apical granules only slightly larger than surrounding granules. Slightly enlarged granule may represent hallucal granule on both feet. Metatarsal granules not present in holotype. Hair on foot covers heel. Terminal pads o f digits weakly striate (Fig. 4).

Ear. Pinna very small, curled external edge on large supratagus. Fawn hairs on postcro-intcrnal and ventral margins o f pinna.

Dentition (Fig. 1). Upper incisors: I1 narrow, peg-like, procumbent, slightly curved, taller- crowned than all upper incisors; 11 separated by large diastema from I2. Left and right 11 widely separated. Crown height, length subequal in I2 and I3.14 slightly greater crown height and length than


I2 and I3. All upper incisors lack buccal cingula yet no lack o f differentiation between root and crown. I4 with no anterior or posterior cusp. Root o f I4 narrow.

Upper canines: C 1 long, slender, caniniform with indistinct boundary between root and crown. No buccal or lingual cingula. No anterior or posterior cusp.

Upper prcmolars: Small diastema between crowns C* and P*, P1 and P2, P2 and M2. P1 shorter crown height than P2. Minute anterior, posterior cingula cusps P 1 and P2. P2 broad postero-lingually.

Upper molars: Posterior tip P' not contacting parastylar corner M2. Anterior cingulum below combined stylar cusp B-paraconc short, broad, complete to prominent paraconulc at base o f paraconc-stylar cusp B apex. Paracone-stylar cusp B o f M2 approx, half height o f mctaconc. Stylar cusp C, E not visible on L or RM2. Trigon basin plunges steeply from prominent metaconule down to weak posterior cingulum.

M3, stylar cusp B and paraconc separate. Broad anterior cingulum contacts metastylar corner M2, tapers down along base o f paracrista, rises steeply to base o f paraconc apex at prominent paraconule. M3 lacks stylar cusps A, C and E. Stylar cusp D prominent and broad. No posterior cingulum; prominent metaconule as in M2.

M4, anterior cineulum complete, broader, much longer than M ' Paraconule, m etaconule prominent. Stylar cusp D reduced but broad. Stylar cusps C, E absent. Posterior cingulum absent.

M5, broad at anterior cingulum terminates quickly away from metastylar corner M4, posterior cingulum absent. Protocone reduced, narrow. Metastylar corner reduced.

Lower incisors: I t much taller crown height than h- *i* *2» b 〇val *n anterolateral view, I( and I2 gouge-like in occlusal. I2 subequal crown height I3. All lower incisors with prominent lingual cusp. I3 with most conspicuous posterolingual cusp almost as (all as primary incisive cusp. Lower canine rests between posterior cusp and primary incisive cusp *3-

Lower canines: Q large, broad, caniniform, characterised by upward, forward projection and sickle-like curvature root to crown tip. No buccal, very weak lingual cingulation, no posterior cusp.

Lower premolars: Premolar row very short, Pj, P2 crushed together, overlapping, separate from M2 by small diastema. Lack buccal, lingual

cingulation. Crown height P2 much taller P卜

Prcmolars relatively broad, bulbous, both with weak posterior cusp, P2 with very weak anterior cusp. Bulk o f each premolar mass concentrated posterior to line drawn transversely through middle o f ihe two premolar roots.

Lower molars: All molars broad, bulbous, tightly crushed together. M2 talonid wider than trigonid, anterior cingulum absent. No buccal or lingual cingulum. Paraconid absent. Mctaconid reduced to small bulge o f enamel subequai height 10 minute entoconid. Metacristid greatly reduced, oblique to lon g axis o f dentary; h ypocr islid m ore perpendicular. Cristid obliqua very short, extends from hypoconid to posterior wall o f (rigonid intersecting at point well lingual to point directly below tip o f protoconid. Hypocristid terminates midway between hypoconid, metastylid. Long, low entoconid. From base o f meiaconid posteriorly entoconid forms prominent semicircular bulge o f enamel (occlusal view) on cndoloph. Indistinct posterior cingulum.

M3, talonid slightly wider than (rigonid. Anterior cingulum very p oor ly developed originating lingually in weak parastylid notch into which hypoconu lid tucked. N o buccal cingulum. Narrow, weak posterior cingulum almost obliterated by crushing anterior cingulum o f M4. Paraconid well developed, smallest trigonid cusp. Entoconid well developed, subequal in height to paracon id. C ristid ob liq u a extends from hypoconulid to posterior wall o f trigon id intersecting trigonid at point directly below tip o f protoconid but well buccal 10 metacristid fissure. From base metaconid posteriorly, endoloph follows line oblique and lingual to dentary axis.

M4, trigonid wider than talonid. Prominent parastylid wraps around hypoconulid M3, weak anterior cingulum on M4. No buccal or posterior cingula. Reduced cristid obliqua intersects trigonid at point well lingual to longitudinal vertical midlinc drawn through tip 〇r protoconid but buccal to metacristid fissure. Entoconid M4 worn but was obviously large, making prominent bulge o f enamel on endolph. Rest o f morphology as in M j except that metaconid is more reduced.

M5, (rigonid much wider than talonid. Anterior cingulum as in Posterior cingulum absent. O f three main trigonid cusps metaconid slightly taller than paraconid but both dwarfed by protoconid. Hypoconid M5 talonid much smaller than \14. Between hypoconid and base o f metacristid, cristid obliqua forms low, very weak cresi, which contacts trigonid wall directly below meiacrislid fissure. Evidence o f small entoconid.

VAN DYCK: A NEW DASYURUS ISI

Sku ll (Fig. 1). D. spartacus is unique amongst Dasyurus species in having a high skull and extremely narrow rostrum between the lachrymal canals. Sagittal and nuchal crests are highly developed. Dorsally the rostrum is gently grooved longitudinally by a depression running.along the nasal sutures. One lachrymal foramen occurs on each side o f the orbital rim. The left and right alisphenoid tympanic bullae are widely separated and relatively small. The foramen pseudovale is small and bisected by a (hin bridge o f the alisphenoid. The foramina o f the transverse canal are large and obvious, whereas the cntocarotid foramina are small and obscure. The internal jugular canal foramina are large; the canals are raised and prominent. The posterior lacerate foramina are large and exposed as are the carotid foramina. The premaxillary vacuities extend from the level o f the I4 root back to the level o f the m iddle o f the C 1 root. The very small maxillary vacuities extend from (he level o f the protoconc root o f M2 back to a level slightly posterior to the protocon c root o f M4. Minute palatine vacuities extend from the level o f stylar cusp B on M5.

Variation in paratypes. Dental and cranial. In adult female PM22003 palatine vacuities are large and more numerous than in the holotype. All (eeth are very worn. The anterior cingulation o f M5 is broad and complete 10 the trigon basin. A very wide diastema separates P j and M2. In juvenile female PM22002 R and l.M4"are partially erupied, R and LM5 are unformed. RM2 shows large slylar cusp C. LM> shows bifid en locon id while L and RM5 are pariially erupted. The unworn dentition o f this paraiypc clearly demonstrates the very reduced m etacon id s o f R and LM 2 and the strong development o f entocon ids on M2_5.

In juvenile male PM22001 M445 are unformed. All other teeth arc partially erupted. Stylar cusp C is present on both R and LM^ but not M3. The crown tip o f 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, M5 is unformed. The entoconids o f R and LM2 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. C o lou r 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 furred and appears m ore golden in co lou r than the

holotype o r paratypes. The tail is short-furred, but demonstrates the golden base which is replaced by black fur one third o f the way down the tail toward the tip. In spirit paratype adult male PM22004 Fuscous black hairs appear slightly bleached to dark reddish brown. The partly e\crtcd penis shows the penis appendage observed and m entioned by Woolley and Webb (1977).

Reproduction. Waithman (ncld notes) recorded 7 nipples for PM22003. Assum ing growth rates similar to those recorded for Dasyurus viverrinus (Fleay 1935), juveniles PM2200I and PM22002, collected 17 May 1972 and 26 June 1973 respectively, may have both been born in February.

Habitat. Specimens o f Z2 were collectedin low mixed savannah at three localities: three from Morehead 8041’S* 141039*E (Rg. 6>; one from Mibini 8°50'S, I41°38E; and one from Mari 9°irS, 141°4rE. PM22001 was "brought in by school children from a road pit*(field notes J. Waithman) and PM10400 was killed while raiding a fowl house. Colleccion details for the other specim ens are not available. Collection areas experience a m onsoonal climate where 75% o f annual rainfall (1682 mm for Morehead) falls during a wet season lasting from December 10 May (Paijmans, Blake, Blecker and M cAlpine 1971).

The habitat type, low-mixed savannah (Paijmans et al. Plate 16, Fig. 2, 1971) occurs in Indorodo, G oe and Mibini land systems where the habitat is inundated and waterlogged during the wet season and burned during the dry season. Trees average approximately IS m in height but som e grow as tall as 28 m. Most com m on species include Tristania su av eo lan s, M ela leu ca sym ph oca rp a , Xanthostem on crenulalus, G revillia glau ca and Banksia denlata. Less com m on species include D ep lan ch ea leiraphy lla, M. v irid iflora , E.

flfg.又 Juvenile female (PM22002>.Phoio: J. Waithman.


Fig. 6. Dasyurus spartacus habitat, low-mixed savannah a( Morchcad, Papua New Guinea. Pholo: H. Parnaby.

polycarpa, T. longivalvis. A cacia spp., D illen ia alala, Parinari nonda and Xanthosiem on barassii. A patchy open shrub layer 1.0-3.3 m in height contains A cacia leptocarpa, Choriceras tricorne, M etasiom a polyant hum, Rhodamnia, A cacia simsii, Tim onius d loch id ion , S in oga tysicephala and \ow Pandanus. The ground layer (with a cover o f 80-100%) con ta in s Im p era ia cy lin d rica , Pseedopogonotherum irriians, Germ ainia capilata, Eriachre squarrosa, Schoenus spp., Sch leria and Rhynchospora rubra. N epenthes and Danella are always present and Ulricularia, D rosera and E riocau ion occur in areas o f very p oor drainage (Paijmans et al. 1971).

Elymol〇K>'. The name spartacus em bodies many features shared by this dasyurid with the notorious Thracian gladiator — strength, a tenacious fighting spirit, and the capacity to draw blood.

D IS C U S S IO N

Past reference to (his new dasyurid as D. geo ffro ii has resulted from not only its strong supcrHcial resemblance lo D. g e o ffro ii but a lso from a reasonable expectation that D. g eo ffro ii with its extensive pre-European distribution throughout Australia (Ride 1968, Archer 1979, Johnson and R off 1982) could occur in New Guinea. One notable feature, however, differentiates quolls north and south o f Torres Strait; the condition o f the premolar row. In Australia the genu s D asyu ru s is characterised by a long premolar row with

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 Dasyurus are generally widely spaced, the loss o f P3 indicates considerable prelusive specialisation. It cou ld 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 m ore primitive condition eg. M urexia /owgfcflMc/a/ff, thelow er premolar row may occupy up to 32% o f the entire lower tooth row. In Phascoga le tapoata/a, a slightly more specialised dasyurid exhibiting a trend toward loss o f P3, the value decreases to approx im ately 25%. In Sarcoph ilu s harrisii where P3 has been lost from both upper and lower tooth rows, the percentage o f Icmw premolar r w drops to approximately 17%.

In D. hallucatus, D. viverrinus, D. m aculaius and D. g eo ffro ii this value ranges from approximately 22-23% and suggests a trend o f prcmolar loss then subsequent rostrum and dentary lengthening — a rc\'ersal o f the trend exhibited by Sarcophilus. The greatly reduced premolar rows o f D. spartacus and D. a lbopunciatu s (19-20??) appear to demonstrate the prcmolar condition just subsequent to the loss o f P3 and would suggest that these two species were in Tact more primitive than all the others.

The possibly Pliocene Dasyurus dunm alli from the Chinchilla Sand, with its single rooted P3 could lend support to this argument. With a crushed and on ly slightly greater proportion o f premolar row (22%), D. dunm alli cou ld represent the species ancestral to D. spartacus and D. albopunciatus. However, (he same constraints that Bartholomai (1971) and A rcher (1982) app lied to the interpretation o f the significance o f D. dunm alli in D a syu ru s ph y lo gen y m ust b e app lied s im u ltan eou sly to D. sp a r ta cu s and D. albopunciatus. In these two species a consideration o f character states apart from the prcmolar row indicates specialisation o f a highly derived nature eg. d isproportionate C 1, V-shapcd upper incisor row, metacone M2 perpendicular to stylar cusp D, M2 reduced without entoconid, specialised penis, m etacrista M 3 shorter than in M4, greater reduction o f paracones, metaconids reduced and upper premolars bulbous (see Tabic 3).

These fea⑴res clearly make D. spar/acus and Z). albopunciatu s (along with D. dunm alli for fw e r


reasons) inclligiblc for consideration as the most plesiomorphic members o f the genus when the morphologically primitive D. hallucatus is considered. If D. dunmalli was considered the sister species o f D. spartacus and D. albopunctatus, then loss o f P3 must have occurred once in D. hallucatus and then reversal o f the premolar character state cx:curred once in the D. dunmalli lineage whereupon the persistent Dasyurus 2-premolar trend reemerged in D. albopunctaius and D. spartacus (Fig. 7A). While this is a tempting and parsimonious approach it assumes many synapomorphies from D. dunmalli o f which so little material (a few incomplete dentaries) is available.

Archer (1982) has suggested that loss o f P3 may have occurred twice within the genus, once from the primitive hallucatus-albopunciaius stock (I do not accept the association o f albopunctaius with hallucatus as ancestral in Dasyurus) and then once again from the common ancestor (dunmalli-Wke) o f the species vi’ve/r//i叫容 and moew/a/ui：. However, two synapomorphies (more reduced metaconids and broad, oval premolars) o f dunmalli and the spartacus-albopunctatus-maculatus group suggest that D. dunmalli is more closely related to these three species than to viverrinus and geof/roii. In this case loss o f P3 may have occurred once in the hallucatus-viverrinus-geof/roii line and then once again in (he ancestor (D. dunmalli?) o f D. spartacus and D. albopunctaius.

If consideration is given to Bartholomai's (1971) warning regarding the dubious nature o f speculating on the relationships o f D. dunmalli then another approach (free from the influence o f D. dunmalli) is represented in Fig. 7B. To opt for the most parsimonious approach the assumption in this case is made (hat the crushing o f the premolar row in D. spartacus and D. albopunctatus is a highly derived condition associated with the shortening of the rostrum. Archer (1976) notes that the reduction o f P3 and widening o f the premolar tooth row are derived states within the dasyurids as a whole. Bensley (1903) related the tendency o f 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 vertebrate-killing dasyurids and Thomas (1887) related premolar reduction to specialisation in dasyurids. In this interpretation, D. spartacus and D. albopunctatus are sister species more closely related to one another than either is to D. geof/roii.

The upshot o f these considerations is twofold. Firstly that regardless o f a preferred phylogeny, D.

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Fig. 7. Two cladograms o f the hypoihetical relationships o f Dasyurus spartacus. Cladogram A includes the Pliocene species D. dunmalli and reflects interpretaiion 4A o f Tabic 3. Cladogram B includes only extant members o f ihe genus and rcflccis imcrpreiafion 4B o f Table 3. See Tabic 3 for reference to all other apomorphies depicted in the cladograms.


1. l ' n a r r o w , p e g l i k e P P A P P P -

2 . I - p r o c u m b e n t , s e p a r a t e P P A P P A -

3 . c l p r o p o r t i o n a t e P A A A A k A

A. P r e m o l a r r o w 1 o n g v p r e m o U r s S p d c e d ^ A3i n

A3T T

A3T T

A ? A2T T T T

5 . P r e m o l a r s n a r r o w , s m ^ l l p P P A A A A

6 . M o l a r s n o n - b u l b o u s p P P P P A P

7. I n c i s o r r o w V - s h a p e d p A A A A A -

8 . P a r i c o o e s r e d u c e d A1 A2 A2 A ? A2 A ? -

9 . M e t a e o n i d s u n r e d u c e d P A2 A1 A ? A3 A3 A2

10. H e t a c o n c H ? p e r p . t o S t . c u s p D P A A A A A -

11. H e t a c r i s t a e q u a l t o o r l o n g e r P A A A A A -

12. H - u n r e d u c e d P A A A A A -

13. w i t h m e t a c o n e P A A A A A -

14. w i t h e D t o c o n l d A A A A A P -

15 . S k u l l h e i g h t l o w P P P A P P -

16. B r o a d r o s t r u m w i d t h b e t w e e n l a c h r y m a l s P P P A P P -

17 . P e o n s i m p l e P A A A A A -

18. P o s s e s s i o n o f h a l l u x P P A2 A1 P P -

19. H in d f o o t s h o r t a n d b r o a d P P A P P P •

Table 3. Character states o f features thought to be useful in phylogenetic interpretation o f quoll evolution. Character 4 presents two interpretations (sec discussion), a. (hat the premolar slate in D. dunmalli represents the least derived o f 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.

spartacus is a highly specialised species, more derived than D. geo ffro ii but not as specialised as D. maculalus. Its greatest affinities are with D. albopunctaius. Secondly that D. albopunctaius, far from being the traditionally accepted primitive species so frequently associated with D. hallucalus is one o f the more highly derived members o f the genus.

One could have hoped for a more comfortable phylogenetic scenario from the tropical rainforests o f New Guinea out o f which such a parade o f primitive laxa has emerged in the past.

ACKNOWLEDGEMENTSThe examination o f type material held in overseas

institutions was made possible through the financial support o f the Royal Zoologica l Society o f New South Wales, the Linnean Society o f New South Wales and the Queensland Museum Board o f Trustees. I gratefully acknowledge the help o f the following for allowing me to examine specimens in their care: J. Menzics (National Museum and Art Gallery, Papua New Guinea), L. Gibson (Australian Museum), D. Kitchener (Western Australian Museum), C. Kemper (South Australian Museum),


A. Allison and C. Kishinami (Bernice Bishop Museum, Hawaii), M. Tranier (Museum National D'Histoire Naturelle, Paris), C. Smccnk (Rijksmuseum van Natuurlijkc Histoire, Leiden), P. Jenkins (British Museum, Natural History, London), R. Angcrmann (Museum fur Naturkunde dc Humboldt-Universititat zu Berlin) and G. Musscr (American Museum o f Natural History, New York). 1 also thank H. Parnaby (University o f New South Wales) and J. Waithman (Conservation Commission o f The Northern Territory) for collection details and photos o f specimens, and particularly M. Archer (University o f New South Wales) and R. Raven (Queensland Museum) for their helpful discussions with me. B. Cowell (Queensland Museum) produced the photographs.

REFERENCESArcher, M., 1976. The dasyurid dentition and its

relationships to lhai o f diddphid、 thylacinids, borhyacnids (Marsupicarnivora) and peramelids (Pcramdina: Marsupialia). Ausi. J. Zool. Suppl. Ser. 39: 1-34.

A rcher, M., 1978. The naiure o f the molar-prcmolar boundary in marsupials and a reinterpretation o f the homology o f marsupial cheekteeth. Mem. Q d Mus. 18: 157-164.

Archer, M., 1979. The status o f Australian dasyurids, thylacinids and myrmccobiids. Pp. 29-43 in The status o f endangered Australasian wildlife cd by M. J. Tyicr. Proc. Roy. Zool. Soc. SA: Adelaide.

Archer, M., 1981. Results o f the Archbold Expeditions. No. 104. Systematic revision o f the marsupial dasyurid genus S/n/n/Aopsic Thomas. /tm". Mus. Nai. Hist. 168: 61-224.

Archer, M.. 1982. Revicu- o f (he dasyurid (Marsupialia) fossil record, integration o f data bearing on phylogenetic interpretation, and supragcncric clas&ificaiion. Pp. 397-443 in Carnivorous marsupials cd by M. Archer. Royal Zoological Society o f New South Wales: Sydney.

Arnoi.d, J. M., 1983. Western Quoll Dasyurus geof/roii P. 22 in The Australian Museum com plete book o f Australian Mammals cd by R. Strahan. Angus and Robertson: Sydney.

Bartholomai. A., 1971. t/i/mrifl"/, a newspecies 〇r fossil marsupial (Dasyuridac) in the upper Cainozoic deposits o f Queensland. Mem. Q d Mus. 16: 19-26.

Bensi ev, B. A., 1903. On the evolution o f the Australian Marsupialia; wilh remarks on the relationships o f the

marsupials in general. Trans. Linn. Soc. Lond. (Zool.). 9:83-217.

Fleay, D., 1935. Breeding o f Dasyurus viverrinus and general observations on the species. J. Mamm. 16(1): 10-6.

Honacki, J. H., Kinman, K. E. andKoeppl, J. W., 1982. Mammal species o f the world. Allen Press Inc. and The Association o f Sysiematics Cotlections: Lawrence, Kansas, USA.

Johnson, K. A. and Roff, A. A., 1982. The Western Quoll Dasyurus geof/roii (Dasyuridae, Marsupialia) in (he Northern Territory: hi&iorical records from venerable sources. Pp. 221-226 in Carnivorous marsupials cd by M. Archer. Roy. Zool. S o c New South Wales: Sydney.

Kirsch, J. A. W. and C ai.aby, J. H., 1977. The species 〇r living marsupials: an annotated list. Pp. 9-26 in The b iology o f marsupials ed by B. Stonehouse and D. Gilmore. Macmillan: London.

Kirsch. J. A. W.. and Archer, M., 1982. Polythetic cladistics. or, when parsimony's noi enough: th« relationships o f carnivorous marsupials. Pp. 595-619 in Carnivorous marsupials cd by M. Archer. Royal Zoological Society o f New South Wales: Syxincy.

Paijmans, K., Blake, D. I. T., Blecker, P. and McAlpinf., J. P., 1971. Land resources o f the Morehead-Kiunga Area, Territory o f Papua New Guinea. CSIRO Land Res. Ser. 29: 1-125.

Ride, W. D. L., 1%8. On ihc past, present and future o f Australian mammals. Ausi. J. Sci. 31: 1-11.

R idgway, R.. 1912. C o lo r standards and c o lo r nomenclature. Publ. by author: Washington.

Tate, G. H. H., 1947. On the anatomy and classification o f the Dasyuridac (Marsupialia). Bull. Amer. Mus. Nai. Hist. 88: 97-156.

Tavlor, J. M., 1984. Mammals o f Australia. Oxford University Press: Melbourne.

Thomas, O., 1887. On ihc homologies and succession o f the teeth in ihe Dasyuridae with an attempt to trace the history o f the evolution o f mammalian teeth in general. Philos. Thins. R. Soc. Lond. 178: 443-62.

Waithman, J., 1979. A report on a collection o f mammals from soulhwest Papua, 1972-1973. Ausi. Zool. 20(2): 313-326.

W oo lley , P. and WbBB, S. J., 1977. The penis o f dasyurid marsupials. Pp. 307-23 in The b io logy o f marsupials cd by B. Stonchousc and D. Gilmore. Macmillan: London.

Z iegler, A. C . 1977. Evolution o f N w Guinea's marsupial fauna in response to a forested environment. Pp. 117-138 in The b iology o f marsupials ed by B. Stonchousc and D. Gilmore. Macmillan: London.

Documents

THE BRONZE QUOLL, DASYURUS SPARTACUS (MARSUPIALIA: …bionames.org/bionames-archive/issn/0310-0049/11/145.pdf · spartacus (spirit specimen PM22004) 2.1 mm, D. geoffroii (spirit specimen