Spilocuscus kraemeri ( Schwarz, 1910 )

Pine, Ronald H., Mack, Andrew L. & Timm, Robert M., 2017, Marsupials And Rodents Of The Admiralty Islands, Papua New Guinea, Occasional Papers of the Museum 352, pp. 1-30 : 4-16

Spilocuscus kraemeri ( Schwarz, 1910)

Admiralty Cuscus

The literature on this species is complex, confusing, widely scattered, and in various languages; thus we thought it useful to summarize, analyze, and annotate the published pertinent information on the species. Schwarz (1910) named “ Phalanger maculatus krämeri ” from the “Admiraltäts- und Hermit-Inseln” and on the basis of 35 skins and 7 skulls in the Berlin Museum [given as “ 32 specimens ” by Aplin et al. (2015:50)] and collected by “… THILENIUS, SCHOEDE und … KRÄMER.” The new taxon was said to be most closely related to “ Ph [alanger]. m [aculatus]. papuensis ” [now generally regarded as a full species, Spilocuscus papuensis (Desmarest, 1822) ] “from Waigeoe” (= Waigeo). The male of the new species was described as having a white to yellowish ground color with moderately large blackish-brown spots with faded borders. The snout, cheeks, and forehead were said to be mostly a paler rusty brown; the limbs spotted, (the spots?) somewhat paler than [on?] the back; the tail with large brown spots. The chin and throat white, the chest and belly yellowish. The female was described as unspotted with the snout and forehead reddish brown, the top of the head, nape of the neck, and the back blackish brown, not shiny. The rear part of the back was described as reddish brown, the tail somewhat darker, the limbs reddish brown, the chin white, the throat and upper part of the chest blackish brown, the ventral portion of the chest and belly yellowish, the underside of the base of the tail dark brown. Skull measurements were given for the holotype, an old male, variously identified as “Berl. Mus. A. 12.6.09” and “A. 126 09”, “gesammelt von SCHOEDE;” and for a subadult female, “Berl. Mus. A. 19.07.2.” No definite localities, even to island or island group, were given for any specimen. Although earlier in the description Schwarz had stated that the specimens were from both the Admiralties and the Hermit Islands (p. 406), he later (p. 408) wrote “Hab.: Admiralitäts-Inseln.” The skull and dentition were described and the former compared with those of specimens of Spilocuscus maculatus (sensu lato?) that Schwarz had seen from other localities. Although Schwarz did not indicate a type locality, Feiler (1978) provided a color drawing of an individual identified as Berlin Museum “12609” and from Manus, presumably the holotype.

Thomas (1914) listed four specimens of “ Phalanger maculatus krämeri, Schwarz ” from Manus. Two in the BMNH were collected by Meek and Eichhorn and two were listed as being in the Tring Museum.

Cohn (1914), unaware of Schwarz’s (1910) description of S. kraemeri , redescribed his animal as Phalanger maculatus minor , on the basis of at least 80 specimens from Manus. These specimens presumably are or were housed in Bremen. However, our attempts to confirm this have not met with success. He also stated that over 60 living individuals from Manus had passed through his hands. No type specimen was designated, no catalog numbers or museum(s) of deposition were mentioned, and no localities, other than the names of islands, were given. In addition to Manus, Cohn stated that he had seen this species on Ponam, and on Lou, where he found it to be quite common. He attributed its presence on these islands to the persistence of primary forest, although Flannery (1995b:105) found this species to be “common in all vegetation types, from gardens to almost undisturbed forest” on Manus. Cohn stated that on smaller, more heavily populated islands, the species had been extirpated, but mentioned only Baluan as being one of these islands (see also Helgen and Jackson 2015). It is unclear whether Cohn actually had any evidence of the former occurrence of S. kraemeri on any of these islands or merely surmised that it had once occurred there. The latter seems more likely, although Flannery (1995b) accepted the former presence of S. kraemeri on Baluan, based on Cohn’s statement (see also Helgen and Jackson 2015). Cohn found S. kraemeri to be very common on Manus and to be the most important game animal for the locals. Cohn was on Manus from September until December of 1912. During this time, he examined many pouch young ranging in size down to 95 mm head plus body length and 80 mm tail length. These smallest ones were blind and hairless. Cohn concluded that this species did not breed all year long. He stated that the heavy rains began in November (with the dry season starting around March/April) and it appeared to him that breeding must have ceased a short time before this. He concluded that a second breeding season must exist early in the year, making the species diestrous, because in September he found numerous already half-grown animals. Flannery (1995b), in discussing Cohn’s findings, apparently thought that all of the small September–December young were around the minimum size given and omitted the evidence for a second breeding season. Aplin et al. (2015:56) reported a “fully furred pouch young” during the period 2–6 October 2014. According to Helgen and Jackson (2015:496), “Limited observations [unspecified] suggest the Admiralty Spotted Cuscus breeds seasonally, giving birth generally to a single young in July–August, with pouch young observed in September–December (then carried by mothers on their backs); young achieve independence in June.”

According to Cohn, the females are more numerous and larger than the males. The males were described as having dark brown legs with little white on them, the belly white with very small brown spots, the tail as a rusty-brown. In these respects, Cohn’s description differed somewhat from that of Schwarz (1910). Two male specimens out of considerably more than 20 were a uniform silver gray with small dark dots on the back; the base of the tail with a faint brownish wash. Both of these males were half grown and were taken to Cohn at the same time and he speculated that they might have been from the same litter.

Cabrera (1919) synonymized Phalanger maculatus krämeri Schwarz with Phalanger maculatus minor Cohn and placed [ Phalanger ] “ minor y fusca ” Oken, 1821 in the synonymy of Phalanger orientalis (Pallas, 1766) . In this, he seems to have been following Thomas (1888:201), who had placed “Phalangista minor and fusca (Oken) Schinz, Cuv. Thierr. … (1821)” in “ Phalanger orientalis .”

According to Schwarz (1934:90), in the course of his idiosyncratic, essentialistic speculations concerning the phylogeny of and also supposedly directly environmentally-induced color patterns in cuscuses, certain spotted patterns belong “to the segmental type, being homologous to the rosettes of a leopard, and representative of the ‘English’ type [undefined] of piebald pattern … these spots evolve a tendency to spread, thus in the end producing a true piebald pattern, such as is found in the males of Ph. maculatus . [new paragraph] That the very distinct markings found in this species are derived from the ones just described is shown by the females of Ph. m. papuensis Desmarest from Waigen and of Ph. m. krämeri Schwarz from the Admiralty Islands, some of which develop spots, but of a less progressive type than is found in the males, and more resembling that of the male Ph. o. ornatus.” Schwarz (1934:90–91) went on to write “Males which are nearly or perfectly white are … found in Ph. maculatus , although in that species perfect albinism is rare, and a stage similar to the ‘Siamese’ or ‘Marten’ type of cats and rabbits is the rule … the individuals show the normal piebald pattern earlier in life.”

Tate and Archbold (1937:381) noted that “ Minor Cohn from Admiralty Islands would appear to be a synonym of krämeri Schwarz …” and stated that the Archbold collection held topotypes of “ krämeri.” They figured the skull, minus the mandible, of AMNH 99900, seen from below. Concerning “ krämeri ” they wrote (p. 383) that the pelage “… appears to be thoroughly and consistently distinct from that of our Arfak and south New Guinea specimens … A striking characteristic of krämeri in both sexes is the nearly black head on which the cinnamon-colored ears stand out contrastingly, and the deep chestnut-colored fore limbs.” The Archbold holdings were listed (p. 384) as “Admiralty Islands (Manus Isl.): 1 y. ad. ♂; 3 y. ad. ♀ s (all except one ♀ skins only); 3 y. ♂ s, 1 y. ♀ (skulls only); 1 ad. albino ♀.” Concerning the last, Tate and Archbold wrote (p. 384) “The muzzle and the insertions of the superciliary and labial vibrissae are reddish brown … and the skin and hairs of the hands and feet are yellowish ….” The supposed “albino” is presumably AMNH 99749 , which was recorded as having a brown iris, apparently the normal color—the collector, W. F. Coultas, also recorded a brown iris for specimens with normally colored pelage. Our notes on this specimen state “All white dorsally except some very faint buffy areas on rump and base of tail. Ventrally, base of tail and adjacent area faint buffy. Some buffy toward end of tail. Grayish around pouch area and posterior to that but maybe soiled.” In their tables of measurements (pp. 448–449), the following specimens are listed: “A.M. 99832 juv. ♂ … Manus”, “A.M. 99831 juv. ♂ … Manus”, “A.M. 99829 y. ad. ♀ … Admiralty Is.”, “A.M. 99900 y. ad. ♀ … Admiralty Is.”, “A.M. 99749 ad. ♀ (albino) … Admiralty Is.”

Tate (1945:27) mentioned “Our [AMNH] original small series of skins and skulls from Manus … including one white female.” Tate, who treated this species as “ Phalanger maculatus krämeri ,” mentioned the Echiquier (Ninigo) Islands as included in its range but provided no further information. Tate also, somewhat tentatively, synonymyzed Phalanger maculatus rufoniger Zimara, 1937 [= Spilocuscus rufoniger (Zimara) ] of mainland New Guinea, with S. kraemeri . He (p. 19) characterized S. kraemeri (including rufoniger ) as “Male with dark head, most spots dark, on white ground, female dark throughout, with reddish tail, hands, and feet. A white aberration occurs.” Concerning the Ninigo Islands, Jentink (1885:117–118) had reported “ Cuscus maculatus ” from there on the basis of “Young male, Museum Godefroy, Hamburgh, 1869. Collected by Captain Alfred Tetens. Head, ears, hands, feet and spots on tail and on outerside of legs red; spots on back and nape of neck brownish red …Adult female. Museum Godefroy, 1869. Collected by Captain Alfred Tetens. Tail injured. Crown of head, nape of neck, back and flanks sooty red; face and legs red; ears and tail yellowish red; underparts like in other females.”

Laurie and Hill (1954) used the name Phalanger maculatus kraemeri for this form and, presumably following Tate (1945), treated it as not even subspecifically distinct from Spilocuscus rufoniger and listed it as occurring in the Echiquier (Ninigo) Islands and on the Huon Peninsula. They placed Balantia minor Oken, 1816 in the synonymy of Phalanger orientalis (Pallas) .

Haltenorth (1958:28) wrote, concerning this animal, “Huon-F[leckenkuskus]., P. m. krämeri Schwarz, 1910; Huon-Halbinsel, Kaiser Wilhelms-Land, Admiralitäts- u. Echiquier-Inseln.”

Collins (1973:177) gave the range of “ Phalanger maculatus ” as including “the Ninigo and Admiralty Islands, New Britain , New Ireland, and some of the Bismarck Archipelago.”

Kisokau (1974) stated that he had collected “ Phalanger maculata ” during the period 22 December 1971 to 19 January 1972 on “Rambutso Island” [= Rambutyo Island] and Lou. No information concerning where the specimens were deposited was given but it seems likely that it was in a collection that was housed in the old Department of Environment holdings at Moitaka, much of which was lost to mold, decay, and general lack of curation by the 1990s (information provided by an anonymous reviewer). All subsequent authors except for Menzies and Pernetta (1986) seem to have overlooked Kisokau’s publication.

Feiler (1978) examined 32 skins and 2 skulls from Manus. These were some of the 35 skins and 7 skulls that had been the basis for Schwarz’s original description of the species. Feiler gave the type locality as “Admiralitäts-Inseln.” He mentioned one small white individual (“gelblichweiss oben und unten”) which he thought was a young animal but said that the remainder were spotted. Feiler made it seem that the three females mentioned by Schwarz (1910), and showing the unspotted female pattern, were unavailable for some reason. Schwarz did not mention a white specimen. Feiler stated incorrectly that Schwarz had written that S. kraemeri ’s range reached to eastern New Guinea (the island). Feiler also mentioned two female, Berlin Museum, skins damaged by the war (“ZMB, 11. 9. 1899 ”), collected by Thilenius, presumably from the Admiralties, that he said that Schwarz must have seen. Feiler then divided the males into four groups on the basis of color pattern. For some reason, he listed only 30 of the 32 specimens he had examined. His first group consisted of the single whitish specimen. His second was individuals with the palest [or most pronounced?] spots, with the snout rusty red, rust-colored to reddish– black; the front feet and nape of the neck more or less paler or darker rusty (from golden yellow to reddish– brown); with only isolated white spots; pale rusty–red spots on the back, isolated darker spots between them; about 60% of its surface white, tail almost solidly rusty red with isolated pale spots.

The third group had the face until above the eyes rusty red, the neck rusty to almost black and without spots; the forefeet darker, reaching, in two cases, a blackish reddish brown, in general like in his second group. The ground color cream with pale golden reddish brown tinge, 60–80% of the surface of the back is white with large spots; the tail reddish brown more or less paler or darker and with white spots. Feiler mentioned one specimen from “ 12. 6. 1909 ” (recall that the catalog number of the holotype was variously given by Schwarz as “A. 12.6.09” and “A. 126 09”) and taken by Schoede, identified to locality only by “Deutsch-Neuguinea,” as having the ground coloration more yellow and with the dorsal spots not reddish–black but a deep and sharply contrasting black with 80% of its surface white.

The fourth group was made up of animals with dark skins, the forefeet a dark rusty red, large spots, with 30–50% of the skin white, intensely black spots on a yellow–white background, sometimes with isolated reddish spots in between, the tail reddish above, a pale yellowish–white underneath, the hind end above and below the same, one individual from “D.-Neuguinea” with the tail underneath also mostly reddish brown. A skin from “Hermit-Inseln,” “Berlin Museum 5. 7. 1909,” taken by Krämer resembled the specimens from Manus, the nape of the neck was very rusty, which is seldom the case in specimens from Manus. The tail was said to be reddish brown as in the specimens from Manus.

Feiler discussed two skulls from what he called the “Hermit-Inseln,” one from “Echiquier,” “MNHL” (apparently a typo for MNHG, meaning from the Museo Civico di Storia Naturale “Giacomo Doria,” Genoa) taken by Godeffroy, and a specimen in the Berlin Museum, “Agoma [?] A 19. 07.” taken by Krämer.

Koopman (1979) referred to animals from theAdmiralty and Ninigo Islands as P [halanger]. maculatus .

George (1987) treated S. kraemeri as belonging in the species S. maculatus (É. Geoffroy St.-Hilaire) and, tentatively, as constituting a recognizable subspecies.

Flannery and Calaby (1987) discussed one young female (AM 3693) and two male specimens (AM 3563, 5117). AM 5117 was labeled as from New Britain (which Flannery and Calaby thought was an extremely unlikely provenance) and the other two were Taronga Zoo (in Mosman, New South Wales) specimens of unknown origin. These specimens were identified as of S. kraemeri because they “conform closely to the morphology of kraemeri as described by Tate (1945) [see Tate’s brief statement quoted above].” In addition to there being these Australian zoo specimens, the Field Museum in Chicago houses a male specimen FMNH 74914 ( Figs. 1A , 2 ), represented by a skin and skull, from the Brookfield Zoo and labeled as from Australia, that clearly represents S. kraemeri . The history of this specimen may be tied in with that of the Australian Taronga Zoo specimens examined by Flannery and Calaby (1987). Paradiso and Fisher (1972) reported, for example, the importation of eight live “ Phalanger maculatus ” into the U.S. in 1970. Collins (1973) also discussed S. maculatus (including S. kraemeri ?) being held in the Taronga Zoo and in at least one U.S. zoo. FMNH 7494 is discussed further in this species account under “Specimens examined.” Pine has seen a specimen (apparently of S. maculatus sensu stricto) from a U.S. pet store and now housed in the Los Angeles County Museum.

Flannery and Calaby (1987) also mentioned a specimen from Mussau Island in the St. Matthias group as possibly belonging to S. kraemeri , but Flannery and White (1991), Flannery (1995b), Helgen and Flannery (2004), and Helgen and Jackson (2015) reported only S. maculatus from there. This last publication’s treatment of the cuscuses on Mussau is odd, considering that Helgen (2007) treated the cuscus on Mussau as belonging to an unnamed introduced species.

Flannery and Calaby (1987:555) wrote “… we consider it extremely unlikely that kraemeri reached Manus with the aid of man, as it is a highly distinctive form, and given the lack of distinctive species to have evolved on islands isolated since the end of the Pleistocene in Australia, it appears to have been isolated on Manus for a considerably longer period.” Nonetheless, Flannery and Calaby continued to treat kraemeri as a subspecies of S. maculatus . They noted (p. 555) that “Two color morphs have been recorded for adult female Spilocuscus from Manus. Feiler (1978) mentions a single white female from Manus. [In fact, Feiler indicated that the specimen was of a male— Tate (1945) mentioned a white female.] However, most individuals seem to be very dark-colored, with a large black saddle area on a reddish background.” Although Flannery and Calaby (1987) provided color photographs of a skin of a typically spotted adult male [AM 3563] and a living juvenile male from an unknown locality, and Flannery (1995b) published a color photograph of a living adult and typically spotted male from Manus, we have found no photographs of an animal identified as an adult female. Feiler (1978) and Flannery (1994), however, provided colored drawings of adult males and females. Feiler’s figured female from “Admiralitätsinseln” was Berlin Museum 11. 9. 1899, the male, Berlin Museum 12609 from Manus. Although Flannery (1994:218) stated that “The male [of the species] has small blackish spots on a white background …,” the spots on the male figured do not appear small and the earlier literature reports size of the spots as ranging from small to large. Flannery and Calaby (1987:554) wrote “It is possible that the white females from Manus and Mussau represent a distinct species related to the mainland S. maculatus and that the highly coloured kraemeri represents a separate species.” Singadan (1996), however, assumed that white specimens of both sexes were only color variants in animals of the S. maculatus species group on the mainland of New Guinea, the Admiralties, and “the St. Matthais Islands.” Flannery and Calaby (1987) provided descriptions of the color patterns of the specimens available to them. Their photograph of the juvenile male shows a gray animal with a white cheek. Singadan (1996) wrote that juvenile males are normally gray with grayish brown spots and patches. The two more-or-less adult males described by Flannery and Calaby are said to be (p. 554) with “limbs and tail … brownish–red. The crown … a darker shade of brown … base coloration … white … dark mottling … the same colour as the dark area of the crown over the shoulders … becomes darker, almost black over … lower back. Near … base of … tail … spots lighten again [this variation in darkness of dorsal spots not evident in their published color photo]. Ventrally … white, with brown hairs on … lower sides of … neck.” They described (p. 554) a juvenile female as “… ventral surface has a white patch on the chest with a longer white patch over the belly. A darker line … surrounds these light patches. Overall … dark red- dish–brown, with silvery red guard hairs. On … base of … tail … lighter patch of fur … under fur … lighter coloured.” Although Singadan (p. 78) wrote that the tail in both male and female S. kraemeri is “white with shades of brown or yellow,” the specimens and figures we have seen show warm pale brown or more or less tan tails, although some white may be on the tails of some. Singadan gave body mass as 2.2–3.2 kg. Being unaware of the later archaeological discoveries to be made on Manus, Singadan hypothesized that the Admiralties cucuses had been introduced from the mainland about 3,000 years ago and had acquired their distinguishing characteristics since then.

Menzies (1991) provided a map indicating the range of S. m. kraemeri to consist of Manus and an unidentified dot in the ocean which is probably intended to represent Wuvulu, an island that is questioned as a habitat for S. kraemeri by Flannery (1995b), although it is listed from there by Helgen and Jackson (2015). Menzies gave “Admiralty Islands” as the type locality. He stated (p. 68) that isolated populations of Spilocuscus maculatus occur on “Wuvulu, Manus Islands [sic] but not New Britain. ” Writing of S. maculatus in general and presumably including S. kraemeri, Menzies stated (p. 69) that “Male spotted cuscuses are bigger than females ….” Helgen and Jackson (2015) stated the opposite, as had Jentink (1885), Cohn (1914), and Helgen and Flannery (2004). According to Helgen (2007), females average 14% heavier than males.

Flannery (1995b:104) treated kraemeri as a full species, stating “These species [ kraemeri and maculatus ] differ in dentition and skull features and are probably not closely related.” The animal from Mussau Island was then apparently treated as S. maculatus maculatus . It was unequivocally treated as S. m. maculatus by Flannery (1994), apparently, also by Helgen and Flannery (2004), and definitely by Helgen and Jackson (2015).

Flannery (1994, 1995b) made no mention of the existence of white female individuals. He stated (1995b:105) “… archeological evidence suggests that it [ S. kraemeri ] has reached Manus recently, perhaps in the last one or two thousand years ….” Flannery (1994) and Flannery et al. (1994) had earlier expressed this same view. Helgen and Flannery (2004:831), however, wrote that at “Pamwak, a rockshelter site in southern Manus … S. kraemeri is represented erratically in the record at least as far back as about 11,000 years ago. This … suggests to us that S. kraemeri might be a natural element in the Manus fauna.” In addition to Manus, Flannery (1995b:105) listed Lou and Luf (Hermit Islands) as places where S. kraemeri occurs and “Ninigo” and Wuvulu Island as places where it possibly occurs. Flannery (1994, 1995b) cited Cohn (consistently misspelled “Kohn” in 1994) (1914) as the authority for the species having been extirpated on Baluan and other, unspecified, islands. Groves (2005a) listed it only for Manus and Lou, as had Flannery (1994).

As late as 1988, Flannery (1995b:105) found S. kraemeri to be common on Manus “in all vegetation types, from gardens to almost undisturbed forest.” The IUCN Red List of Threatened Species lists it as “Near Threatened” (see Helgen et al. 2008). Helgen and Jackson (2015:480) wrote that this species “… appears still to be common in all parts of its limited distribution ….”

Flannery (1994, 1995b) gave “Yowat” as the Nali language name (“South Manus ”) for this animal.

The colored drawing of the male, in Flannery’s 1994 book, is based on AM M20925 , from Polomou, Manus. The female depicted was acquired at the Lorengau Market on Manus. The specimen from Polomou is the only one of which we are aware which has had a fairly precise field locality published for it. Flannery (1994) provided measurements from at least two males and two females from Manus. Natural history information is provided by Flannery (1994, 1995b).

Singadan (1996) noted that S. kraemeri is a lowland animal, usually occupying lowland rainforests, usually below 1,000 m. However, the maximum elevation of Manus is only about 720 m and no other island inhabited by this species is even that high. Singadan also noted that there were at least 52 specimens held in the collections of the University of Papua New Guinea and PNGMAG. Assuming that the 80 or so specimens seen by Cohn are still in existence, then the number of available specimens is indeed impressive.

Williams (1999) reported S. kraemeri remains from as far back as ca. 11,000 ybp, at the Pamwak archeological site on Manus.

Anthony (2001) discussed the purchase, in a market in Kimbe, New Britain , of a skin of a cuscus which had reportedly been speared on the small island of Bali off the north coast of New Britain. (Bali is an alternative name for the island generally known as Unea or Uneapa in West New Britain .) Judging from the photograph provided of this skin and our direct examination of the specimen (UWZM M27790 ; Fig. 1B ), it is that of a male S. kraemeri , although Anthony identified it as that of an S. maculatus .

Helgen and Flannery (2004:828) wrote, concerning Spilocuscus wilsoni , that the “adult male coat is pure white dorsally and ventrally … as in some … S. m. maculatus …”, taking no note of white individuals of both sexes, reported of S. kraemeri , although Flannery and Calaby (1987) had earlier mentioned white Spilocuscus from Manus and which they supposed might represent a second species there. [Somewhat incongruously, Helgen and Jackson (2015) in their discussion of S. kraemeri , mention that there are white individuals of S. rufoniger ( Zimara, 1937) , but make no mention of white S. kraemeri .] Helgen and Flannery (2004:830) also stated “Two species of Spilocuscus occur in the Bismarck Archipelago of eastern Papua New Guinea: S. maculatus maculatus , which occurs on the islands of Mussau … and New Ireland; and S. kraemeri of the Admiralty group …”. They stated that the Bismarck S. maculatus apparently had been introduced from the island of New Guinea (see also Flannery and White 1991). Although Flannery et al. (1987) give the presence of a protocone on “M1” (the upper deciduous tooth in their reckoning) as a defining characteristic of Spilocuscus, Helgen and Flannery (2004) stated that the diagnostic protocone was on “P3.” Citing Williams (1999), Helgen and Flannery noted that archeological remains of this species are now known from as far back as 11,000 years ago on Manus, which suggested to them that it “might be a natural element in the Manus fauna.” However, they also suggested that the Admiralties populations might originally have been derived from New Britain , an island which previously had been thought to harbor no Spilocuscus . In support of the idea that S. kraemeri might inhabit New Britain , they cited the specimen (AM M5117) earlier mentioned by Flannery and Calaby (1987) as being labeled as from New Britain , and the specimen figured by Anthony (2001) and supposedly from the New Britain satellite island Bali. See our treatment of Phalanger orientalis for an account of Helgen and Flannery’s ideas concerning their hypothesis as to why S. kraemeri is common on Manus and rare on New Britain (if it occurs there at all). Helgen and Jackson (2015:480) wrote, concerning the presence of S. kraemeri in the Admiralties “… may have been introduced to some or all of these islands, and have originally evolved on New Britain. ”

Helgen and Jackson (2015) listed S. kraemeri from the island of Los Negros, which is separated from Manus by a narrow channel, and from Pak Island in the Admiralties, in addition to the islands previously mentioned herein. They also described the ground color of the back of male S. kraemeri as being chocolate brown with creamy-white blotching or spotting. Based on specimens we have seen, however, this pattern has tended to be reversed and the dark areas often darker than chocolate brown. AMNH 99750, from Rambutyo, however, appears to have a bit more dark coloration dorsally than whitish. Aplin et al. (2015) reported that during the time spent on Mt. Sabomu (2–7 October 2014), one adult female with a fully furred pouch young was captured by a landowner and one other individual was observed sleeping in the crown of a tree (page 56), and they state that S. kraemeri has been recorded from Los Negros. It surely is there or was, but we have found no additional records of this, although AMNH 182070 (see below), a zoo-derived specimen, is labeled as from “Las [sic] Negros.” A facial image of a live adult, presumably a female, was provided by Whitmore et al. (2015:xix). Aplin et al. (2015) report that Wildlife Conservation Society staff and University of Papua New Guinea students are studying the ecology of the Admiralty Cuscus at several localities (possibly only on Manus). They state further that the Spilocuscus from the St. Matthias Group (which includes Mussau) represents an unnamed species, and cited Helgen (2007) as their authority. As noted earlier, this is in contrast with Helgen and Jackson’s (2015) account, which lists the animals from Mussau as representing S. maculatus .

Specimens examined.—FMNH 74914, which was referred to earlier, is represented by a stuffed skin in good condition ( Fig. 1A ) and a skull ( Fig. 2 ) in which the right premaxilla is glued in place and the right maxilla and lachrymal had become loose anteriorly. The animal was an immature male received at Chicago’s Brookfield Zoo on 14 July 1953 and which died in captivity on 16 December 1953. The deciduous teeth are still in place, the third molars were unerupted, and the fourth molars are undeveloped ( Fig. 2 ). After the original skin tag was written in ink, someone added the word “ Australia ” to the tag in light pencil. Greatest length of skull is 75 mm. M1 is 6.9 mm in length. Only one small and nonfunctional tooth, rather than two or three such teeth, is developed on each side of the mandible, between the large incisor and the deciduous tooth. This is a common condition in S. kraemeri (see Cohn 1914; Feiler 1978) and is also the case, unilaterally, in three of the seven AMNH specimens we examined and that were collected by Maa. There can be no doubt that this specimen is of S. kraemeri . The color pattern of the pelage possesses all of the diagnostic features as seen in the most typical males of the species, as described and figured in the literature, and as also possessed by male specimens of S. kraemeri examined by us in the USNM ( Fig. 3 ) and AMNH. This includes the very sharply contrasting dorsal and lateral pattern of whitish ground color and bold black spots, the tawny rump and tail, and the distally dark forelimbs with a chestnut suffusion ( Fig. 1A ).

The incomplete skin, UWZM M27790 ( Fig. 1B ), identified on the tag as being of an adult male, likewise appears to unequivocally belong to S. kraemeri . Also according to the tag, it was “Purchased from market in Kimbe, New Britain. ‘Killed with spear on the Island of Bali’ ” just off the coast of West New Britain. The reddish coloration of the rump and tail is deeper and richer than in FM 74914 and the black markings dorsally and laterally are elongated to form a somewhat striped pattern but this falls well within the range of variation in S. kraemeri (see, for example, Feiler’s 1978 plate 2 depicting S. kraemeri ). Here again, the dark coloration becomes more general anteriorly and what is left of the skin of the forelimbs shows a chestnut suffusion. Unfortunately, the skin of the head, the distal portions of the front limbs, of the hind limbs, and the distalmost portion of the tail, is missing in this specimen. The apparent size of the animal that possessed this skin is consistent with that of adult S. kraemeri .

AMNH 193746–193752 : These seven specimens, from a series also including AMNH 193753 (not seen), were collected on 15 December 1959 at Lorengau (purchased in the market?) by T. C. Maa. They were prepared as fluid-preserved heads only, but the skulls of the seven at least, have now been cleaned and the head skins—in poor condition—saved. Until examined by us, these specimens, unidentified to sex and without field measurements or notes, had been identified as of “ Phalanger orientalis intercastellanus .” They range in age from an immature with deciduous teeth, unerupted (but visible in the cleaned skull) third molars, and undeveloped fourth molars (greatest skull length = 68.8 mm) to an adult (greatest skull length = 83.3) with all adult teeth fully in place ( Fig. 4A, B, C ; Fig. 5A, B, C ). In addition to the individuals of these two age classes, there is another with the third premolars fully erupted. All possess only two upper premolars in each maxilla, except for one which bears, unilaterally, only the third.

Additional specimens seen are: AMNH ♀ skin & skull 166953, “ Admiralty Isl. ,” “Manx Zoo, N.Y. Zoo Soc.;” AMNH ♀ ♂♂♂ skulls only 99829–99832 from “ Manus Isl. ,” collected by W. F. Coultas ; AMNH ♀ skin & skull 99900 from “ Manus, Village Drabui ” (coordinates?), external measurements 450, 400, 73, 25, collected by Coultas ; AMNH ♀ skin & skeleton 182070 from “Las [sic] Negros Island; near Manus,” “Gift: N.Y. Zoological Society,” very large skull 94.2 mm including incisors, sutures tending to be fused ; AMNH ♀♀♀ skins only 99747–99749 from “ Rambutyo Is. ,” collected by Coultas, external measurements 480, 390, 67, 20; 500, 430, 67, 23; and 520, 440, 68, 23, respectively. A latex endocranial cast of AMNH 99749 is deposited at the Field Museum , although the skull is now missing from the AMNH. AMNH ♂ skin & skull 99750 from same locality, same collector, external measurements 478, 384, 68, 21.

In addition to the above specimens, we are fortunate to have examined five specimens in the Papua New Guinea National Museum & Art Gallery. These consist of four adult females and an immature male for which developmental information is available. Also, all five are provided with locality data. Previously, only one locality had ever been published. Our figured color photograph ( Fig. 6 ) of a skin of an adult female is the first such photo to be published either in color or black and white. Whitmore (2015:xix) provided a photograph of a head and part of a forelimb of a living animal, however, which might very well be that of an adult female, perhaps the one mentioned by Aplin et al. (2015:56).

The immature male, PNGMAG 22552 ( Fig. 7 ), was shot in “Tall Lowland Canopy” at Kabuli on 17 November 1974. Total length was 447, head plus body length 239, tail length 208, mass 250 g. The milk premolars are still in place, the first molars the only ones erupted. The coloration of this immature is unlike that of any specimens, of any age, reported previously, being a uniform dark drabby brown dorsally and laterally, creamy midventerally and on the lower jaw. Additional data dealing with measurements and dates of collection will be necessary in order to evaluate Cohn’s (1914) conclusion that this species is diestrous.

The four females ( PNGMAG 22430 , 22550 , 22551 , 22553 ) are all age-ranked as old adults, except for 22553, which is ranked as a young adult. All were shot in “ Lowland Rainforest ” or “ Lowland Canopy ” or “ Tall Lowland Canopy ” at “ Kabuli Base Camp, SW Manus I.” or at “ Kabuli (2°7′50″S, 146°40′0″ or 10″E or 2.1305S, 146.6695E).” Measurements of the three old adults are: condylobasal length 82.1–90; total length (for 2) 880, 915; head plus body length 462–498; tail length 417–425; hind foot without claw 58–60; ear from notch 22–26; mass 2.8–3.2 kg. None of the dimensions of the young female exceeded the greatest ones for the older animals and some were smaller than them all. Its mass was 2.2 kg. Coloration in these females was more or less similar. The dorsum anterior to the rump ranged from dark chocolate brown sometimes suffused with chestnut, especially anteriorly and on the crown. The rump and tail fur are contrastingly bright pale golden brown. The legs tend toward chestnut. The cheeks and lower jaw and a midventral stripe are cream-colored, with this stripe sometimes bordered with black or nearblack (see Fig. 6 ).

Eight specimens, all from 1944, were briefly examined at the ( U.S.) National Museum of Natural History. These were USNM 276975–2769781 , 277034 . USNM 276975 was taken on “Ponam Islet.” The rest were labeled as from “ Lorengau on Manus (purchased in market?), except for 277034, labeled as from “Admiralty Islands.” The specimen from “Ponam Islet,” represented by a skin and skull, had a very large, robust skull with the supraorbital region much elevated, as is typical for Spilocuscus maculatus . Only one of the anterior minute lower premolars was present on each side. USNM female 276976 and 277634, provided with both skins and skulls, had skins with color patterns similar to those of females in PNGMAG (see Fig. 6 ), although no sex determination was provided for 277034. USNM female 276977 was cataloged as a skin plus skull specimen but only the skull was found. Both initial upper premolars were absent. USNM males 276978, 276999, 276981, skins with skulls, had pelage pattern and coloration (see Fig. 3 ) similar to that of FMNH 74914 (see Fig. 1A ). USNM 276979 had only one of the minute initial lower premolars present on each side. USNM 276981 lacked the initial right upper premolar. USNM 276980 , an immature, supposedly a skin plus skull specimen, had only the skull at hand. USNM 277003 , a female immature in fluid, was not examined.

The Australian Museum ( Sydney , New South Wales) has 21 specimens of S. kraemeri that were obtained from markets on Manus and 6 specimens obtained from the Taronga Park Zoo.

Figures

Figure 1. Skins of male Spilocuscus kraemeri. A—Stuffed skin of immature male (FMNH 74914), specific locality unknown. B—Incomplete skin of adult male (UWZM M27790) said to be from the island of Bali, off the north coast of New Britain (see text for additional details).

Figure 2. Dorsal, ventral, and lateral views of the skull of an immature male Spilocuscus kraemeri, specific locality unknown (FMNH 74914—see Fig. 1A). Condyloincisive length of skull = 75 mm.

Figure 3. Stuffed skins of adult male Spilocuscus kraemeri from the series of six collected in 1944 at Lorengau, Manus Island (purchased at the market?) by D. H. Johnson. A—USNM 276981; B—USNM 276978; C—USNM 276979.

Figure 4. Dorsal, ventral, and lateral views of crania of Spilocuscus kraemeri. A—AMNH 193746, condyloincisive length of skull = 65.0 mm; B—193748, condyloincisive length of skull = 76.4 mm; C—193747, condyloincisive length of skull = 78.9 mm. Specimens from a series of eight collected in 1959 at Lorengau (purchased in the market?) by T. C. Maa.

Figure 6. Skin of adult female Spilocuscus kraemeri (PNGMAG 22550) from Kabuli, Manus Island. Total length = 892 mm; mass = 2.8 kg; condylobasal length = 82.1 mm.

Figure 7. Skin of an immature male Spilocuscus kraemeri (PNGMAG 22552) obtained at Kabuli, Manus Island, on 17 November 1974. Total length = 447 mm.

Figure. 5. Dorsal and lateral views of the corresponding mandibles from crania in Figure 4 of Spilocuscus kraemeri from the series of eight collected in 1959 at Lorengau, Manus (purchased in the market?) by T. C. Maa. A—AMNH 193746; B—193748; C—193747.