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The Discovery and Early Natural History of the Eastern Pygmy-Possum, Cercartetus nanus (Geoffroy and Desmarest, 1817) PDF

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The Discovery and Early Natural History of the Eastern Pygmy- Possum, Cercartetus nanus (Geoffroy and Desmarest, 1817) Jamie Mark Harris School ofEnvironmental Science and Management, NSW Southern Cross University, Lismore 2480 ([email protected]) HARRIS, J.M. (2006). The discovery and earlynatural history ofthe easternpygmy-possum, Cercartetus nanus. ProceedingsoftheLinneanSocietyofNewSouth Wales 127, 107-124. Early accounts of the eastern pygmy-possum, Cercartetus nanus (Marsupialia: Burramyidae), are reviewedandthehistoryofitsdiscoveryisreported. Fran9oisPerondiscoveredthespecieswhenonashort stay on Maria Island in 1802. Various names have been conferred upon it, but C nanus is now accepted. C The early natural history literature on nanus has some very interesting andhighly relevant accounts of morphology, distribution, behaviour, habitat and diet. Some discrepancies and misinterpretations in the early literature are identified, and several interesting 19* Century illustrations ofC. nanus are reproduced. This study documents the significance of the primary soiu^ce material pertaining to this small elusive marsupial. Manuscriptreceived4May2005, accepted forpublication21 September2005. KEYWORDS: Burramyidae, Cercartetusnanus, discovery, naturalhistory, nomenclature INTRODUCTION CercartetusnanussharesthefamilyBurramyidae withfourotherextantspecies: the long-tailedpygmy- The eastern pygmy-possum, Cercartetus nanus, possum, C. caudatus, little pygmy-possum, C. is broadly distributed in Tasmania and along the lepidus, western pygmy-possum, C. concinnus and eastern seaboard of mainland Australia from south- mountain pygmy-possum, Burramysparvus (Strahan eastern Queensland, through coastal New South 1995). Thispaperinvestigatesthediscoveryandearly Wales and Victoria, and into south-eastern South accountsofthenaturalhistoryofC. nanus,whichwas Australia (Strahan 1995). Currently there are two the first of the burramyids to be formally described recognised subspecies: C. nanus nanus in Tasmania; by Europeans (Desmarest, 1817). Subsequently, and C. n. unicolor on the mainland (Wakefield C. concinnus (Gould, 1845) was recognised, then 1963; McKay 1988). It is a small (~24g) and agile C. caudatus (Milne-Edwards, 1877), C. lepidus tree-dwelling marsupial that feeds chiefly on nectar, (Thomas, 1888) and5./?arvM5 Broom, 1896. pollen and invertebrates within a range of habitats includingheathland, woodland, sclerophyll forestand rainforest. Modem studies have documented some MATERIALSAND METHODS aspects ofthe population biology ofthis species and it is understood that it depends on the presence of a The work of Thomas (1888) is instructive diverserangeoffloweringplants(particularlyBanksia for early accounts of Cercartetus spp., and in this in certain areas), and that seasonal food availability regard 36 references for C. nanus (and its synonyms) influences both the timing and duration ofbreeding were provided from literature published from 1817 (Turner 1984, 1985; Ward 1990; Turner and Ward to 1875. The Kinetica and Firstsearch databases 1995; Bladon et al. 2002). During winter, C. nanus is were used to identify libraries within Australia and ableto storeup fatinitsbodyandtail, andcanexhibit overseas that held the relevant early natural history torpor (Geiser 1993; Turner and Ward 1995; Bladon titles from which copies of the relevant articles et al. 2002). Pygmy-possums have a prehensile tail, were obtained. I also supplemented these papers which resembles that of a ringtail possum, and also by searching for mention ofthe species in the early syndactylous hind feet and an opposable clawless volumes (<1970) ofthe Australian Zoologist and the hallux (Turner and McKay 1989). VictorianNaturalist (Harris 2005). The literature was EARLY NATURAL HISTORY OF CERCARTETUSNANUS examinedandreviewedforinformationon discovery, 1817, 1820; Cuvier 1826; Lesson 1827, 1838, taxonomy, morphology, distribution, abundance, diet, 1842; Temminck 1827; Fischer 1829; Schinz 1844; habitat andbehaviour. Iredale and Troughton 1934; Tate 1945; Wakefield and Wameke 1963). The type specimens for both C. nanus and A. minimus were collected by Peron, but HISTORICAL RECORDS thelatterisconsideredtohavecome fromWaterhouse Island, which lies close to the north-eastem coast of Discovery Tasmania (Wakefield andWameke 1963; Rounsevell ThefirstspecimenofC.nanusknowntoEuropeans 1989). was collected by Fran9ois Peron, a naturalist aboard NicolasBaudin'svoyagetothesouthseas ontheships Taxonomy and nomenclature Le Geographe and Le Naturaliste (1800-1804). His Upon the retum of the Baudin expedition to discoveries and observations whilst inAustralia have France in 1804, several ofthe greatFrenchzoologists long interestedhistorians (Triebel 1948; Faivre 1953; of the period, including Anselme-Gaetan Desmarest Cornell 1965; Plomley 1983; Wallace 1984; Homer and Etieime Geoffroy Saint-Hilaire worked rapidly 1987; Plomley et al. 1990; Hunt 1999; Anderson describing and classifying the specimens collected 2001). He is credited with the collection of about by Peron. In the encyclopedic Nouveau Dictionairie 100,000 zoological specimens, 2500 of which were d'Histoire Naturelle, Desmarest (1817) described new to science, including C. nanus. Whilst on a short the small marsupial collected from Maria Island as stay on Maria Island, off eastern Tasmania between Phalangista nana Geoff. (=Geoffroy). However, 19 and 27 February 1802, Peron traded with the subsequently there has been uncertainty as to Aboriginal inhabitants (the Tyreddeme people; Ryan whom the specific name nana ('dwarf') should 1981) for a single small marsupial. Peron (1809:233) be attributed to, with some authors allocating it to wrote (in translation) 'In the class of mammiferous Geoffroy (e.g. Cuvier 1827; Temminck 1827; Lesson animals, I only saw one kind ofDasyurus, which was 1828, 1830 1838; Fischer 1829; Gray 1841; Schinz scarcely as large as a mouse. I obtained one that was 1844; Waterhouse 1846; Gunn 1852) and others alive, inexchange forafewtrifles, fi-om a savagewho to Desmarest (e.g. Giebel 1859; Lydekker 1896; was just going to kill and eat it'. In an unpublished Lucas 1897; Le Souefand Burrell 1926; Iredale and manuscript (now held in the Le Havre Museum in Troughton 1934; Wakefield 1963). McKay (1988) France) Peron also wrote that the animal 'was given statedthatitmustbedatedfromDesmarest[andhence to me by the natives; it was still alive; I believe it to not Geoffroy] as 'Geoffroy's (1803) manuscript was be a new species and have described it as Didelphis never published'. However, Julien-Laferriere (1994) muroides because of its resemblance to the D. mus stated that the species is not mentioned in Geoffroy's ofLirmaeus' (Observations zoologiques by Fran9ois (1803) Catalogue des Mammiferes, contrary to what Peron, on Maria Island, unpublished manuscript McKay (1988) allows to be assumed. Furthermore, # 18043:31). The specimen collected by Peron (a the specimen did not arrive in France until 1804. juvenile male) was transported back to France, and Although Geoffroy did not write on the species, is now held in the Museum National d'Historie Beaufort (1966) believed that Desmarest's allocation Naturelle in Paris as the holotype (Julien-Laferriere ofthe name to his colleague was intentional (also see .1994). Cercartetus nanus still presumably inhabits Desmarest 1820), and accordingly he proposedthat it Maria Island, as there is a relatively recent record should officially be attributed to both as Cercartetus from 1969, when two young animals were found in a nanus (GeoffroyandDesmarest, 1817). Inthis, Ihave dead tree being cut for firewood (Animals and Plants followed Beaufort (1966). Protection Board 1969). In a new edition of Nouveau Dictionairie Plomley et al. (1990) erroneously stated that the d'HistoireNaturelle, published in 1818, a description single small marsupial collected on Maria Island by of P. nana equivalent to Desmarest (1817) was Peronwasthetype specimenforAntechinusminimus. also published. This is significant because the This was probably based on a similar mistake made 1818 edition is sometimes incorrectly referred to by Waterhouse (1846) which was highlighted by as the first description for the subject species (e.g. Wakefield and Wameke (1963). Waterhouse (1846) by fredale and Troughton 1934; Marlow 1962; interpreted Peron's statement offinding a 'Dasyurus' Wakefield 1963; Green 1974; McKay 1988; Tumer as meaning that the dasyurid A. minimus was also and McKay 1989; Flannery 1994; Menkhorst 1995; collectedfromMariaIsland,whenevidently C. nanus Tumer and Ward 1995). Following Desmarest was the only mammal species collected (Desmarest (1817), briefdescriptions ofthe species appeared in 108 Proc. Linn. Soc. N.S.W., 127, 2006 HARRIS J.M. '%-i^-: >-^.__ ...-<r„ Figure 1.ThisillustrationaboveoftwoPhalangistagliriformis{=Cercartetusnanus)appearedinanarticle byThomasBellpublishedintheTransactionsoftheLinneanSocietyofLondonin 1829.Theanimalsappear tobequitelargeduetothedisproportionallysmalltreetrunkandbranches uponwhichtheyarestanding. Proc. Linn. Soc. N.S.W., 127, 2006 109 EARLYNATURAL HISTORY OF CERCARTETUSNANUS \f'r,ff'' a/ ^mm^ , ^ ^ ^ Figure 2. Pouch and extremities of Phalangista gliriformis {=Cercartetus nanus) by Bell (1829). a. Pouch and teats, shortly after the period of suckling; b. Pouch and teats of the unimpregnat- ed animal; c. Prehensile extremity of the tail; d. Fore-foot, upper part; e. Fore-foot, under part; f. Hind-foot, upper part; g. Hind-foot, under part; h. Curl of the tail, observed during sleep. 110 Proc. Linn. Soc. N.S.W., 127, 2006 ) J.M. HAI^RIS subsequent zoological publications printed in French common brush-tail possum. According to Wakefield (Desmarest 1820; Cuvier 1826; Lesson 1827; 1828; (1963), the reference was drawn from anunpublished 1830; 1838; Temminck 1827; Fischer 1829), English manuscript written by Constantin Gloger, but when (Cuvier 1827) and German (Schinz 1844) and were the work was published in May 1841, the name either taken from the original reference or from the Cercaertuswasnotmentioned.Instead,Gloger(1841 specimen which formed the subject ofit. proposed the quite different name PsilogrammUrus On 4 November 1828, Thomas Bell read before for P. vulpina, and used Cercartetus for P. nana. the Lirmean Society of London a description of a Cercartetus makes some reference to the tail (from supposed new species of Phalangista, which he the Greek kerkos) but the significance is obscure named P. gliriformis (Bell 1829). The species name (Strahan 1981). It is not known whether Burmesiter was derived from the latin word 'glires' meaning (1837) incorrectly cited Gloger (unpublished) or if 'dormouse'. His address was based on close substantial changes were made to the work prior to examinationoftwolivefemaleswhichwere 'received publication. Perhaps due to the conftision, the name from New Holland' (Australia), but from what part Cercartetuswas atthattime basically disregarded for was not stated. Bell (1829) detailed a great deal of P. nana. However, itis clearthatthename Cercaertus carefiil observation, but he failed to persuasively is a junior synonym of Trichosurus and not of distinguish P. gliriformis from P. nana. According to Cercartetus (Iredale and Troughton 1934; Wakefield the description, the distinction was proposedbecause 1963; McKay 1988). ofslight differences in the colouring, and principally In a report dated 10 July 1841, and published in because fur was absent from the ears. Bell's November of that year, Dr I.E. Gray of the British coirfidence in the distinction relied on the phrase 'les MuseumsetoutareviewoflocalitydataonAustralian oreilles sont arrondies et couvertes de polls' from mammals wherein he proposed the genus Dromicia Temminck's (1827) description of P. nana, which for P. nana because 'the dentition and the peculiar quoted Desmarest (1817), and translates as 'the ears form and character ofthe tail ofthis species, at once are round and covered with hair'. Later, Waterhouse pointoutthatitshouldconstituteadistinctgenusfrom (1841)statedthat'Temminckshouldhavesaidthatthe the otherPhalangers, fromwhich itdiffers inmanyof ears are covered with very minute hairs, for so small its habits' (Gray 1841). This was later acceptedbyDr are they that to the naked eye they appearnaked' (see G.R. Waterhouse ofthe BritishMuseum (Waterhouse alsoWagner 1843). TheholotypeofP. nanacontained 1846), and subsequently the name D. nana was in the Paris Museum, and also the type specimen widely applied, although the synonym 'Phalangista for P. gliriformis, were re-examined by Waterhouse nana' persisted in a small number of articles (e.g. (1841) and no specific differences were perceived Gunn 1852; Gulliver 1875). Cobbold (1868) reported by him (see also Waterhouse 1846; Wagner 1855). that Professor Richard Owen, of the Royal College Despite this taxonomic oversight. Bell's observations of Surgeons London, disagreed with Gray (1841) on living specimens resulted in some very interesting on the justification of Dromicia. Owen stated that notations on the habits of the species and he also 'modifications oftheteethareunaccompaniedbyany provided some remarkable illusfrations (reproduced change of general structure or ofhabit, whilst those as Figs 1 and 2). However, one inaccuracy in Fig. 1 teeth which most influence the diet are constant' (loweranimal)istheinclusionofaclawonthehallux. and also that 'these differences of dentition are Itshouldbehighlightedthataverysimilarillustration unimportant, and afford no grounds for subgeneric to Fig. 1 appeared in Cobbold (1868), but the hind distinctions'. However, in this case at least, Owen's feet were also drawn incorrectly (see reproduction of view didnot gather support. this image and comments in Strahan 1981). The species was not found on the Australian Thereissomeconftisionintheliteratureregarding mainland until Gerrard Rrefft of the Australian a statement made by Burmeister (1837) which Museum made a report ofa Dromicia found near St. translates as 'a specific genus (Cercaertus Glog.) is Leonards, North Shore, Sydney, New South Wales. formed by the common brush tailed Ph. vulpina\ Krefft (1863) believed it represented a new species It has occasionally been presumed that Cercaertus anddescribeditasD. unicolor, whichwas areference was a mis-spelling or synonym of Cercartetus (e.g to its uniform mouse-colour. However, M.R. Simpson 1945; Marlow 1958; HickmanandHickman Oldfield Thomas ofthe British Museum doubted the 1960; Sharman 1961; Bartholomew and Hudson significance of the find, and believed that Krefft's 1962; Grzimek 1975). In fact, the name Cercaertus Dromicia was probably a D. nana from Tasmania was used in reference to Phalangista vulpina, which which had escaped from captivity (Thomas 1888). isanabsolute synonymfor Trichosurusvulpecula,the He argued that apart from Krefft's specimen, the Proc. Linn. Soc. N.S.W., 127, 2006 111 EARLY NATURAL HISTORY OF CERCARTETUSNANUS species had never been recorded from the mainland, Forbes-Leith and Lucas 1884) or from Tasmania also adding the questionable statement that it is 'to (as accepted by Iredale and Troughton 1934). He be found in the collection of almost every dealer in compared the teeth of nanus (Desmarest 1817), live animals'. Thomas (1888) also remarked that he gliriformis (Bell 1829), unicolor (Kreffl 1863) and had inspected drawings of the premolars of the D. britta (Wood Jones 1925), but could not resolve nana held in the Paris Museum, and compared these the matter with the specimens available to him. with Bell's D. gliriformis. He concluded they were Nonetheless, he suggested that the subspecies should synonymous, which supported Waterhouse's (1841) be C. nanus nanus forTasmania; C. nanusgliriformis earlier view, although Thomas did not mention (=unicolor) for New South Wales and Victoria, and Waterhouse in relation to this. C. nanus britta for South Ausfralia, which was at In 1925, Frederic Wood Jones, Professor of variance from Iredale and Troughton (1934). Tate's AnatomyattheUniversityofAdelaide,communicated (1945) proposal was not adopted because he failed some observations in the Transactions of the Royal to demonstrate unequivocally that gliriformis was Society ofSouthAustralia on what he believedwas a from the mainland. However, Iredale and Troughton newspeciesofDromicia(WoodJones 1925).Anadult (1934) had not proved that Bell's specimens were male, collected at Millicent in south-eastern South Tasmanian. Australia, was described as the type of Dromicia Thenextimportantcontributiononthetaxonomy britta. Certain measurements were provided which of C. nanus was a review by Norman Wakefield of suggestedthathis specimenwas considerably smaller Monash University, who discussed the distribution, than Krefft's D. unicolor and the average specimens habitat and taxonomy ofthis species and the pygmy- of D. nana. For this reason, and also because his possums more broadly (Wakefield 1963). He revised specimen had a greyer colouration, and shorter tail the taxonomy insofar as reducing the number of than D. nana. Wood Jones (1925) believed that it subspecies advanced by Iredale and Troughton should be given species status. It is worth noting that (1934) from three to two, because he believed that measurements for two D. nana individuals were also on the mainland there was only one subspecies, presented by Wood-Jones (1925), but it is, apparent which was reasonably uniform and continuous in thatthesestatisticsareinerrorsincetheyrepresentdata distribution from South Australia through Victoria frommore thantwo animals (seeThomas 1888). This and into New South Wales (see also Le Souef and inaccuracy may or may not have influenced Iredale Burrell 1918). That is, Wakefield (1963) accepted C. and Troughton (1934) to reject the proposed specific n. unicolor as the mainland subspecies, and made C. distinction, but britta was nevertheless recognisedby n. britta an equivalent synonym, while also accepting them at the subspecific level (see below). C. n. nanus as the Tasmanian subspecies. However, The genusnameDromiciaGrayhadbeenapplied in a subsequent note, Wakefield (1970) questioned for close to a century when Iredale and Troughton his own sub-specific assignment, stating that the four * (1934) noted that Cercartetus Gloger antedated cranial specimens available to him from Tasmania Dromicia by several months. They advanced the were 'insufficient to demonstrate difference from or name Cercartetus nanus to supersedeD. nana, which affinity with' mainland populations. Despite this, the included a change in the ending ofthe specific name arrangement of Wakefield (1963) has been in place from nana to nanus to accord with the gender ofthe for more than 40 years (McKay 1988; Turner and new genus (Strahan 1981). Iredale and Troughton Ward 1995; vanWeenen2002), andthis is despite the (1934) then somewhat arbitrarily accepted three sub- absence of any review, testing or elaboration upon species: (1) C. nanusnanusforTasmania,with/! nana which to substantiate this hypothesis. and P. gliriformis as synonyms; (2) C. nanus britta Confiision is even greater in vernacular for south-eastern South Australia with D. britta as a nomenclature. Names included dwarf phalanger synonym; and (3) C. nanus unicolor for New South (Desmarest 1817; Cuvier 1926; 1827), minute Wales and Victoria withD. unicolor as a synonym. phalanger (Waterhouse 1838), dwarf cuscus (Gloger From the type of C. nanus held in the Paris 1841), pigmy phalanger (Waterhouse 1841), Bell's Museum, G.H.H Tate of the American Museum of Dromicia{Gray 1843;Gerrard 1862),opossummouse Natural History, had the skull extracted and cleaned (Gunn 1852; Bonwick 1858; Lord and Scott 1924; for study in 1937 (Tate 1945). He examined the Tate 1945), duskyDromicia, pygmy opossum (Kreffi dentition ofthis and other specimens in London and 1864), thick-tailed Dromicia (Krefft 1868; 1871; sought to determine whether the type ofgliriformis Le Souef 1907), mouse-like phalanger (Cobbold was from mainland Australia (as implied by several 1868), common dormouse-phalanger (Thomas 1888; authors subsequent to Bell 1829, e.g. Gould 1863; Lydekker 1896), dormouse phalanger (Waterhouse 112 Proc. Linn. Soc. N.S.W., 127, 2006 HARRIS J.M. 1846; Lucas 1890; Le Souef and Burrell 1926; soft short hair, ofa whitish colour, Marlow 1958), common dormouse-opossum (Ogilby and are fiimishedwith fourrows of 1892); dormouse possum (Brazenor 1950), pigmy long black vibrissae, the posterior opossum (Le SouefandBurrell 1918),pigmypossum ones tipped with light brown. The (Iredale and Troughton 1934; Wakefield 1963) and eyes are very large, remarkably eastern pigmy possum (Ride 1970). A standard name prominent, and of a jet-black finallyeventuatedwhenacommitteeoftheAustralian colour: the ears of considerable Mammal Society recommended 'eastern pygmy- size, erect, totally destitute ofhair, possum' in 1980 (Strahan 1980). and ofa uniform mouse-colour'. Dentition and Morphology Intermsofcolouration,thefiirwasfirstdescribed Desmarest (1817) stated that the teeth, as far as as grey lightly frosted with a reddish tinge and white it was possible to observe them on this little animal, underneath (Desmarest 1817) and more simply appeared to be arranged like those of phalangers. as upper parts grey, but white underneath (Cuvier Similarly, Bell (1829) stated that the incisors did 1826; Lesson 1827; Schinz 1844; Krefft 1871). In resemble other species of the genus Phalangista, characteristic detail. Bell (1829) stated that his living but complained of the difficulty of examining the examples were: minuscule teeth on living subjects. Owen (1845) 'covered with a very soft and pointedoutthatthespecies 'hasonlythreetruemolars thick fur; the hairs which compose on each side of the jaw', and also that 'the last and itbeingofagraycolourtippedwith penultimate premolars on the lower jaw are. shaped reddishbrown, givethegeneralhue like canines'. Subsequently, Krefft (1863, 1864) was ofrufous-gray. The under parts are able to provide the following dental formula: more sparingly covered with fur of a pale yellowish-gray colour, the M 13-3/1-1 C 1-1/1-1 P 3-3/3-3 3-3/3-3 yellow predominating at the sides, Total = 36 and especially at the throat. The general colour of the face is also The basic phalangerid dentition is three yellowish, the upper and back part premolars and four molars in each row (Tate of the head assuming the rufous- 1945), although Cercartetus is unusual in having gray colour ofthe back'. only three molars in each row, and C. nanus has a diagnostic P^which is large and double-rooted (see Bell (1829) also noted a blackish ring around also Smith 1971; TumbuU and Schram 1973; Green the eye, and remarked on 'a darkish ring partially and Rainbird 1983; Menkhorst and Knight 2001). surroundingtheearsattheanteriorpart,interruptedby In terms ofmorphology, Desmarest (1817) made a distinctwhite spotbehind each (ear)'. Krefft (1863) adescription fi-om a spirit specimen andbrieflynoted described the fiir as 'a uniform mouse-colour lighter it as the size of a mouse, and with a brown circle onthesidesandbeneath,withablackishpatchinfi-ont around the eyes, and imprecisely described the ears of the eye'. Gould (1863) stated that 'considerable as short, rounded and 'covered with hair'. As already diversity of colour exists in different individuals; in mentioned, it should have been stated that the ears some the upper surface is nearly uniform grey, while appear nearly naked. A more articulate description inothers afine tawnyorrufoustintpervades the same was provided by Bell (1829) who stated that: parts; andexamplesareconstantlymetwithexhibiting 'the general form of this every variety of intermediate shade'. Wakefield animal resembles that of the (1963)pointedoutthattheTasmanianmembersofthe common dormouse; but it is larger, species (C. n. nanus) 'have a warm brown infiision broader and more depressed. The in the general body colour and are yellowish on the head is broad across the ears, fi-om sides andunderneath', while the mainland form (C.«. whence it tapers to the nose, which unicolor) 'is less brown and less yellow' (see also Le is somewhat pointed. The nostrils SouefandBurrell 1918). are narrow, and of a semicircular Earlynaturalistsnotedthat C. nanushave several form: the upper jaw, which is features in common with other possums, such as the elongated, overhangs the imder, prehensiletailandfeetspeciallyadaptedforclimbing. and almost entirely conceals it. They also noticed the incrassated base of the tail, The lips are scantily covered with and considered this to be a unique and characteristic Proc. Linn. Soc. N.S.W., 127, 2006 113 EARLYNATURAL HISTORY OF CERCARTETUSNANUS PLATE XVII. "X? COMMON DOfi.MOUSE-PHAl.ANGER Figure 3: This illustration ofthe Common Dormouse Phalanger {=Cercartetus nanus) appeared in Lydekker's Handbook to the Marsupialia and Monotremata in 1896. attribute of this species (Bell 1829; Lesson 1830; Bell (1829) noted that two toes on each of Gray 1841; Waterhouse 1846; Le Souef and Burrell the hind feet were 'united together' (Fig. 1). This 1926). Lydekker (1896) noted the tail as 'rather long morphological feature (syndactyly) is an adaptation with the basal inch thickened', but the incrassation for fur cleansing (Ride 1978) and for an arboreal was not evident in the illustration he provided which lifestyle (Hall 1987). Krefft (1863) noticed that the was originally published in Waterhouse (1841) (Fig. tongue is 'fiimished with a slight brush at the tip', 3). Le Souefand Burrell (1926) explained that 'when and he interpreted this as an adaptation for nectar- capturedin summerthe tail is notusually incrassated, feeding. Thomas (1888) noticed that there were five and the animal is slender and mouse-like; but as large pads on each ofthe palms and soles. There are winter approaches it becomes bulkier, the base of various other minor descriptions of morphological the tail becomes very swollen, and the appearance features outlined in the early literature, but I have of the animal is very much changed' (see also Le only coveredthose ofmost significance. Souef and Burrell 1918). An assessment of the female reproductive organs by Bell (1829) revealed Distribution and abundance fourteats, andmany subsequentnaturalists concurred In the early years of European settlement of with this observation (Lesson 1830; Wagner 1843; Australia it was presumed that the species was Giebel 1859; Thomas 1888; Ogilby 1892; Le Souef peculiar to Maria Island and mainland Tasmania and Burrell 1926; Troughton 1943; Wakefield 1963). (Cuvier 1827; Waterhouse 1838; Gray 1841; 1842; However, in more recent times Wakefield (1970) Gunn 1852; Gould 1845; Waterhouse 1846; Gervais reported an individual with five nipples, and Turner 1955; Giebel 1859; Cobbold 1868). It is now clear (1981) found that there are actually six teats, four that the species also has a broad distribution in the developed and two rudimentary. coastal regions of south-eastern mainland Australia 114 Proc. Linn. Soc. N.S.W., 127, 2006 HARRIS J.M. (Turner and Ward 1995). In the early years however, responded to Waite (1904) with a convincing list the specimens which reached the British Natural of reliable mainland records. Further relatively History Museum were mainly Tasmanian (Gray early (<1970) locality records for Victoria include 1843; Gerrard 1862; Thomas 1888; Wakefield 1963) Heathcote, Blacks Spur, Sale, Avoca, Buanger, which probably led Gould (1863) to postulate that Portland, Erica, Wilson's Promontory, Mount Lock, the species was 'abundant ...in Van Dieman's Land Tamboon Inlet, Mallacoota, Whitlands, Nowa Nowa, (=Tasmania), particularly the northern parts of the Snake Valley, Rushworth Forest, Cape Conran, island'. Lord and Scott (1924) also suggested that it Grenville, Yackandandah and Mount Drummond was more common in northern Tasmania. However, (Harris2005).Acomprehensivereviewofmorerecent by the early 1960s it was considered that the species Victorian records is given by Harris and Goldingay was rare in this State because of 'marked changes (2005). NSW in vegetation' brought about by periodic forest fires Early C. nanus records fi-om include those (Wakefield 1963). Important early literature records fi-om St. Leonards in 1863 and Jindabyne in 1903, for Tasmania include Hobart, Waratah, Launceston, Fitzroy Falls in 1914, La Perouse prior to 1918, Westbury district, and Fury Gorge near Cradle Royal National Park in 1925 and Bowral in 1939 Mountain, Cloudy Bay, Mount Wellington (see (Le Souefand Burrell 1918; Wakefield 1963). Krefift Wakefield 1963), and also Maria, Bruny, Flinders, (1864) stated that 'the range ofthis species probably King and Cape Barren Islands (Le Souef 1929; does not extend beyond the east coast districts' but Hickman and Hickman 1960; Wakefield 1963; Green qualified this by noting that because it is diminutive 1969; Green and McGarvie 1971; Whinray 1971; and nocturnal 'it will be a difficult task to obtain Hope 1973). More recent Tasmanian records and a many examples, and so define its geographical comprehensive distribution map are provided by distribution with certainty'. As further information Munks et al. (2004). became available, Marlow (1958) was able to state NSW While C. nanus was apparently not found on the that its range in was 'between the Hastings mainlandpriorto 1854(Seebeck1995),themaincredit River and Sydney' and extended west only to the for its discovery on the continent should go to Krefft Blue Mountains. Subsequently, Wakefield (1963) (1863), who collected a specimen at St. Leonards, a remarked thatNewcastle was the northern limit ofits suburbofSydney,NSW. However, itis acknowledged range. However, a recent review of the distribution NSW that Bonwick (1858) had earlier noted that 'opossum of C. nanus in (Bowen and Goldingay 2000) NSW mice' occurred at Warmambool, Victoria, but no indicates that its range in extends to Grafton, NSW specimen was collected. The first collected specimen Maclean andTweed Heads and onthe farnorth fi^om Victoria appears to have come fi-om Western coast, althoughmost records are from the south coast Port in 1880 (Wakefield 1963), and subsequently and on the eastern side ofthe Great Dividing Range. A Forbes-Leith and Lucas (1884) accepted the species few scattered western records have been identified as a component of the Victorian mammalian fauna. for Pilliga, Coonabarabran, Dubbo, Parkes and Othervery earlyVictorian records include specimens Molong. The scarcity ofrecentrecords inBowen and collected fi-om Gerabrook and Muckleford in 1886, Goldingay (2000) has ledto its currentrecognition as and Mordialloc in 1887 (Wakefield 1963). Thomas a 'Vulnerable' species inNSW. (1888) was evidently unaware of these Victorian South Australia (SA) and Queensland form recordswhenhedismissedKrefft's(1863)observation the western and northern limit, respectively, of the ofthe mainland occurrence ofthe species. distribution of C. nanus. There are only a small In 1896, Dr Robert Broom recorded that he number ofrecords from each ofthese States. Wood found a large number ofteeth and upper jaws of C. Jones (1925) reported that the first SA specimen was nanus in a sub-fossil bone breccia deposit near the discovered at Millicent, and this specimen is now Wombeyan Caves (Broom 1896). In the same year, held in the collection of the British Natural History Professor Baldwin Spencer of the University of Museum (Wakefield 1963). Only three specimens Melbourne provided details of several specimens from this State were acquiredbythe SouthAustralian secured in southern Victoria (Spencer 1896). Museum prior to 1997, and its status was considered Surprisingly however, its natural occurrence on the rare. These records are confined to the far south- mainland was still disputed. Waite (1904) provided east ofSA. An intensive survey ofthis region which details of a specimen collected at Jindabyne, NSW, targeted C. nanus in 1997 produced a fiirther 27 but was reluctant nonetheless, to declare that the records, and subsequently the status of C. nanus in species definitely occurrednaturally onthe continent. SA was changed to 'Vulnerable' (under Schedule 8 Hall (1904) finally put the controversy to rest, and of the National Parks and Wildlife Act 1972) (van Proc. Linn. Soc. N.S.W., 127, 2006 115 EARLYNATURAL HISTORY OF CERCARTETUSNANUS Weenen 2002; Carthew 2004). In Queensland, the and Burrell (1918) found nests of this species in species was first discovered by Molly O'Reilly in hollow limbs of Eucalyptus squamosa, E. piperata Lamington National Park in 1936 (O'Reilly 1941). andE. haemastoma. Later, these zoologists remarked Furtherexampleswerelaterfoundinthe samegeneral that 'they live in any convenient nook or cranny in a vicinity (Fleay 1966; Wakefield 1970), but as faras is tree, but usually in a hollow limb protected fi'om the known,therangeofC. nanus extends onlymarginally weather, making their nest at an angle. The nest is into Queensland, where it is at present paradoxically composed ofsoft bark, which the animals sometimes rated as 'Common' (Eyre 2004; Harris et al. inprep). have to travel a considerable distance to procure' (Le Souef and Burrell 1926). They also detailed an Diet and habitat observationthatin one case 'itwas aquarterofamile Bell's (1829) captive C. nanus (housed in (~400m) to the nearest tree on which bark similar to London) fed 'on nuts and other similar food'. that in the nest [of C. nanus] was found'. Nesting Captive animals are known to accept arange offoods observationsarescant,butthosepublishedincludethe including bread, cake, seed, honey, milk, cream, discovery ofC. nanusnesting inthe decaying stumps biscuits, lollies, finits and insects (Lord and Scott of grass trees Xanthorrhoea spp. (Green 1969), and 1924; Le Souef and Burrell 1926; Troughton 1931; also in deserted bird and bat nests (Chaffer 1930a,b; Rocking 1939; Conway 1939; HickmanandHickman Schulz 2000). Lord and Scott (1924) commented 1960). In the wild, the first feeding observation was that 'Searching for the retreats of these animals is made by Kreffl (1863) who saw C. nanus 'feeding a tedious task', and that most sightings are 'fi^om on the blossoms of the Banksiae\ He later wrote bushmen who come across them when felling and that 'they live principally on honey and soft insects' cutting up trees in the bush'. They also added that (Krefft 1867). Gould(1863) statedthattheyfeedupon their habits 'naturally make them difficult to obtain, the tender buds and spikes offlowers, which Ogilby and it is more by accident than design that specimens (1892) and Lucas and LeSouef (1909) interpreted are secured'. as meaning that C. nanus was phytophagous. This possum is now generally regarded as omnivorous Behaviour (McKay 1988; Menkhorst 1995; Menkhorst and Bell(1829)wasinpossessionoflivingexamples, Knight 2001), but not herbivorous, and microscopic andthis furnishedhimwiththe opportunityto closely analysis offaeces supports the contentionthatarange observethehabitsofthespecieswhileinconfinement. of dietary items (particularly pollen and insects) are He observedthat: consumed (Huang etal. 1986; Dickman andHappold 'in their habits they are 1988; Tulloch 2004). extremely like the dormouse, As early as 1863 it was recognised that 'of all feeding on nuts and other similar trees itprefersbanksias'(Gould 1863), anobservation food, which they hold in their fore which is supported by modem ecological studies paws, using them as hands [see (Turner 1985; Ward 1990). Bowen and Goldingay also Fig. 1]. They are nocturnal, (2000) and Harris and Goldingay (2005) also note its remaining asleep during the penchantforBanksiahabitat.Earlynaturalistsreported whole of the day, or, if disturbed, that 'they inhabited open wooded country', usually not easily roused to a state of among banksias as well as eucalypts, angophora, activity; and coming forth late in grevilleas, melaleucas and other small flowering the evening, and then assuming shrubs (Le Souefand Burrell 1926; Chaffer 1930a,b). their natural rapid and vivacious While it has been recorded from both wet and dry habits. They run about a small tree sclerophyll forests (Marlow 1958; Green 1973; Harris which is placed in their cage, using and Goldingay 2005), it has been suggested that dry theirpaws to hold by the branches, forests are preferred over wet forests (Wakefield and assisting themselves by their 1963). However, there are both historic and more prehensile tail, which is always recent evidence thatwet forests/rainforestisprobably held in readiness to support them, favouredhabitat onthe edges ofitsrange inTasmania especially when in a descending (Green 1973; Munks et al. 2004) and in Queensland attitude. Sometimes the tail is (O'Reilly 1941; Bowen and Goldingay 2000; Harris thrown in a reversed direction, et al. inprep). turned over the back; and at other Alittleinformationisavailablefi'omtheliterature times,whentheweatheriscold, itis about the nesting requirements ofC. nanus. Le Souef rolled closelyup towards the under 116 Proc. Linn. Soc. N.S.W., 127, 2006

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