Coccymys shawmayeri (Hinton, 1943)

Voss, Musser Scientific Publications Of The American Museum Of Natural History American Museum Novitates Bulletin Of The American Museum Of Natural History Anthropological Papers Of The American Museum Of Natural History Publications Committee Robert S., Meng, Chair Board Of Editors Jin, Prendini, Paleontology Lorenzo, Voss, Invertebrate Zoology Robert S., Whiteley, Vertebrate Zoology Peter M., Knight, Anthropology Managing Editor Mary, Lunde, And, New Guinea Coccymys, Melomys & Bulletin, Albidens Amnh, 2009, Systematic Reviews Of New Guinea Coccymys And ‘‘ Melomys’ ’ Albidens (Muridae, Murinae) With Descriptions Of New Taxa, Bulletin of the American Museum of Natural History 2009 (329), pp. 1-139 : 1-139

Coccymys shawmayeri

7. Sandaun Province, Telefomin Valley, between Telefomin (05 ° 089S, 141 ° 389E) and Lake Louise (05 ° 149S, 141 ° 249E). Lake Louise, westnorthwest of Telefomin, 2800 m: BBM-NG 98492, 100147. ‘‘ NE of Telefomin, ’’ 8000 ft (2440 m): BBM-NG 98365, 98407. Bafunmin (05 ° 069S, 141 ° 269E) and 5 km south, 2300 m: BBM-NG 108076, 108166, 108233. Upper Sol River valley, 2600 m: AM M16745. Flannery and Seri (1990: 190) identified this last specimen as C. ruemmleri , and Flannery portrayed the same animal in a color plate published in his Mammals of New Guinea (Flannery, 1995: 275; T. Flannery, in litt., 2008). Its warm brown fur over the upperparts of the body and and relatively long tail with a long white tip are typical of C. shawmayeri .

8. Sandaun Province, Oksapmin (05 ° 159S, 142 ° 149E), 1850 m: BBM-NG 55461.

9. Enga Province, Kaijende Highlands, Porgera Area (05 ° 289S, 143 ° 119E for Porgera): Menzies (1990: 136) examined specimens (identified as ruemmleri ) from ‘‘Porgera’’ and Helgen (2007b: 60) reported remains (as ruemmleri ) that ‘‘were found in owl pellets in alpine grassland along Waile Creek Road near the Porgera Reservoir [3010 m].’’

10. Southern Highlands Province, Tari region (05 ° 549S, 142 ° 579E for Tari). Tari Gap, 2600 m, BBM-NG 105507, 105534, 105577. Doma Peaks, 2400 m, BBM-NG 105712, 105714. Fringe of Lavani Valley, 2450 m, BBM-NG 60907.

11. Southern Highlands Province, Mt. Giluwe region. 5 km W of Mendi, 2000 m: BBM-NG 60320. Kagaba (05 ° 519S, 143 ° 469E) and vicinity, at base of Mt. Giluwe, 40 road km NE of Mendi, 2800–3600 m: BBM-NG 60524, 60625, 97059, 97157, 97161, 97180, 97238A, 97238B, 97238C, 97241, 97258, 97274A, 97274B, 97280, 97298A, 97298B. North slope of Mt. Giluwe, 3300 m and 3500 m: BBM-NG 101844, 101890. Southeast slopes of Mt. Giluwe, 3600 m: BBM-NG 101924. Southsoutheast slopes of Mt. Giluwe, 2900 m: BBM-NG 101991, 101992, 102008, 102056.

12. Western Highlands Province, Mt. Hagen region. Mt. Hagen (05 ° 459S, 144 ° 029E) 7000–8500 ft (2135–2592 m): AMNH 156475, 156486, 156490, 156513, 156514, 156540, 156546–548, 156588, 156589, 156605, 156664, 156665, 156671, 156673. Tomba, southwestern spur of the Hagen Range (05 ° 509S, 144 ° 029E), 8000 ft (2440 m): BMNH 50.1760, 50.1761. Yanka, eastern slopes of the Hagen Range, BMNH 50.1759 (the examples from Tomba and Yanka were identified as Rattus shawmayeri by Laurie, 1952: 305). Murmur Pass (05 ° 459S, 143 ° 569E), 10–16 km NNE of Tambul, 2700 m: BBM-NG 60685, 60753, 97420, 97456, 97477A, 97556.

13. Western Highlands Province, Nondugl (05 ° 529S, 144 ° 439E): 1600 m, AMNH 222232, 222239 ; 6000 ft (1830 m), AMNH 183609, 183634, 183635.

14. Simbu (Chimbu) Province, Bismarck Range. East slopes of Mt. Wilhelm (05 ° 409S, 145 ° 059E), Lake Aunde, 3570 m (AMNH 192092–97, 192100–102, 192105–107, 192109); 2 mi E Lake Aunde, 3400 m (AMNH 192098); Pengagl Creek 2770 m (AMNH 192103, 192307, 192735); Mt. Wilhelm, ‘‘E. slopes’’ (AMNH 192104, 192113, 192733, 192734, 192736–40); ‘‘Mt. Wilhelm’’ (AMNH 192108, 192114, 192115); Mt. Wilhelm, ‘‘ 9,000 ft’’ (AMNH 156618); Mt. Wilhelm, ‘‘ 9,000 –12,000 ft’’ (AMNH 156608). Consult Brass (1964) for descriptions of the camps at Lake Aunde and Pengagl Creek where some of these AMNH specimens were obtained. We identified an additional 56 individuals (AMNH 276640– 95) that are represented by cranial fragments and dentaries extracted from owl pellets found at 3660 m beneath a boulder on the side of a ridge rising sharply beyond the Lake Aunde camp (Van Deusen’s notes in AMNH archives). High slopes of Mt. Wilhelm: BMNH 50.1763, 50.1764. Mt. Wilhelm, 11,400 ft (3477 m): BBM-NG 100473–77, 100518, 100524, 100557, 100577, 100587, 100619, 100635, 100636, 100644, 100645, 100653–55, 100672, 100673, 100676, 100681, 100696, 100697, 100709. Yandara, Bismarck Range: BMNH 50.1758. Bogo, south slopes of Bismarck Range: BMNH 50.1762 (the BMNH specimens were identified as Rattus shawmayeri by Laurie, 1952: 305). Baiyanka, Purari-Ramu Divide, southeastern part of the Bismarck Range (05 ° 469S, 145 ° 109E) 8000 ft (2440 m): BMNH 1947.1155 (holotype of Rattus shawmayeri Hinton, 1943 ). Mt. Kerigomna (05 ° 589S, 145 ° 079E), 3400 m: BBM-NG 55607.

15. Eastern Highlands Province, eastern end of the Bismarck Range. South slopes of Mt. Otto, Collin’s Sawmill (05 ° 599S, 145 ° 259E), 2300 m: AMNH 192116 ( Brass, 1964, described and mapped this locality). Fatima River, 5 km W of Collin’s Sawmill, 2500 m: BBM-NG 55563, 55582.

16. Morobe Province, Kuper Range, Southern slopes of Mt. Missim (07 ° 149S, 146 ° 509E), 2000 m: one specimen reported (as ruemmleri ) by Willett et al. (1989).

17. Morobe Province, Kuper Range, Mt. Kaindi region. Mt. Kaindi (07 ° 219S, 146 ° 439E): BBM-NG 105375 (2200 m), 51129 (2300 m), 53512 (2350 m), 97718 (8000 ft [2440 m]). Mt. Kaindi Road, vicinity of Wau, 1900 m: BBM-NG 97657. Edie Creek, Mt. Kaindi: USNM 357491.

18. Morobe Province, Wau area. Bulldog Road, ‘‘ 12 mi S’’ of Edie Creek (07 ° 319S, 146 ° 409E), 2500 m: BBM-NG 101267, 101502. Bulldog Road, ‘‘ 12 mi from’’ Edie Creek (07 ° 319S, 146 ° 409E), 2400 m: BBM-NG 61669, 96646, 96802. Bulldog Road, ‘‘32 road km S Wau’’ (07 ° 319S, 146 ° 409E), 2400 m: BBM-NG 29140.

19. Central Province, SSW of Mt. St. Mary (08 ° 109S, 146 ° 589E), 3000 m: BBM-NG 96863, 96864, 96895–97, 96909, 96913 Coccymys kirrhos , n. sp.

20. Morobe Province, Wau area, Bulldog Road, ‘‘ 12 mi from’’ Edie Creek (07 ° 319S, 146 ° 409E), 2500 m: BBM-NG 55824.

21. Central Province, Smith’s Gap (08 ° 039S, 146 ° 539E), ‘‘in the vicinity of Guari,’’ 2500 m: BBM-NG 96950. Kris Helgen wrote us that ‘‘From my understanding, Smith’s Gap is a locality in the border area between Central and Northern Province and is situated at 2500 m. Taylor et al. (1982) gave the elevation of Smith’s Gap as 762 m [2500 ft], which is not correct, and in doing so provided the (artificially) lowest record of occurrence of Rattus niobe , which has been cited since.’’

22. Milne Bay Province, eastern region of the Owen Stanley Ranges, Maneau Range (also spelled ‘‘ Maneao’ ’), north slopes of Mt. Dayman (top camp; 09 ° 499 S, 149 ° 169E), 2230 m: AMNH 158173 (2300 m is noted on the skin tag of this specimen, but 2230 was written in Hobart Van Deusen’s field catalog), 158174,158175 (holotype of Coccymys kirrhos ). The locality was described and mapped by Brass (1956; also see fig. 38) .

23. Milne Bay Province, southeastern ramparts of Mt. Dayman on the Garatin Ridge in the Agaun Region of the Maneau Range , Dumae Creek , 2.3 km N, 0.4 km W of Agaun (09 ° 539S, 149 ° 239E; Dumae Creek , the base camp for the expedition recounted by Cole et al. [1997], is below Garitin Pass , on the road from Agaun to Bonenau ), 1525 m: BBM-NG 109165 .

24. Milne Bay Province, Maneau Range, North slopes of Mt. Simpson, 0.5 km S of Bunisi Village (10 ° 019S, 149 ° 359E), ‘‘Camp 5,’’ 1490 m: BBM-NG 184493.

Brassomys , n. gen., albidens

INDONESIA, PAPUA PROVINCE

1. Northern ramparts of the Snow Mountains, Kelangurr Cave (04 ° 019S, 138 ° 089E), 2950 m, in a valley confluent with the valley of the West Baliem River (see map and description in Flannery, 1999): AMF 134074, 134096, 134126. These three specimens are dentary fragments extracted from

Pleistocene sediments excavated at Kelangurr Cave and were identified by Musser.

2. Snow Mountains, Lake Habbema (04 ° 499S, 138 ° 419E), 3225 m: AMNH 150821 (holotype of Melomys albidens ).

3. Snow Mountains, Bele River valley, 9 km northeast of Lake Habbema (04 ° 059S, 138 ° 509E), 2800 m: AMNH 150531, 150541, 150607, 150618, 150923.

CHARACTERISTICS OF COCCYMYS

The type species of Coccymys is ruemmleri 1, described as a species of Pogonomelomys by Tate and Archbold (1941: 6) who, under the heading ‘‘General Characters,’’ characterized it as a ‘‘small dark brown rat with Melomys -like dentition, feet unmodified for climbing… but with dorsal 3 cm. of tail provided with tactile surface for prehension.’’ They followed with this description:

Skin with rather long fur (12 mm.), colored above near Bone Brown, becoming Clove

Brown along the back, the face with a grayish cast; underparts grayish white, the long pelage with fuscous bases which show through. Hands and feet clothed with light yellow-brown hairs.

Ears fuscous. Tail brown; the scales small and lacking the prominent keeled structure of true

Melomys , their scale-hairs blackish, 3 per scale and from 2 to 3 scale-lengths. … Skull much smaller than that of P. mayeri , the rostrum much compressed and less shortened; supraorbital ridges undeveloped; braincase fuller; zygomatic plate much narrower, and nearly straight as in Macruromys ; palate with posterior foramina; back of palate even with back of M 3;

bullae small; angular process of mandible quite short. Incisors narrow, unbroadened, orange.

Molars very small and of characteristically simple Melomys type.

Tate (1951: 316) was uneasy about the generic allocation of ruemmleri , noting that

1 The name is spelled ‘‘ rümmleri ’’ in Tate and Archbold’s (1941) original description, a patronym honoring Hans Rümmler’s contribution to systematics and evolution of New Guinea rodents ( Rümmler, 1938). Except where we quote authors using the original spelling, we use ‘‘ ruemmleri ,’’ thus conforming to the edict promulgated in Article 35.5.2.1 of the International Code of Zoological Nomenclature (fourth edition, 1999: 40): ‘‘In the case of a diacritic or other mark, the mark concerned is deleted, except that in a name published before 1985 and based upon a German word, the umlaut sign is deleted from a vowel and the letter ‘e’ is to be inserted after that vowel.’’

Pogonomelomys ‘‘remains an unsatisfactory genus due to the fact that it seems to be diphyletic or even triphyletic.’’ He recognized two subdivisions within the genus, the ‘‘ mayeri-bruijnii group,’’ which contains the only two species that now constitute Pogonomelomys ( Menzies, 1990; Musser and Carleton, 2005), and the ‘‘ sevia-rümmleri group.’’ Laurie and Hill (1954: 127) simply listed ruemmleri as a species of Pogonomelomys without question in their list of the land mammals of New Guinea, and before that Ellerman (1949: 88) had suggested a relationship between Pogonomelomys ruemmleri and P. tatei , a form named and described by Hinton (1943).

Tate’s separation of ruemmleri from the other species of Pogonomelomys was later supported by data from study of phallic morphology of endemic New Guinea murines by Lidicker (1968: 641) who noted that it ‘‘is evident from phallic morphology that P. ruemmleri and P. mayeri are not especially closely related, although both fall in the Uromys group. … The latter is close to Melomys , whereas the former is a highly specialized type most closely allied to Uromys and Hyomys . More species of this genus obviously need to be examined, but at least the genus as now constituted seems to be polyphyletic.’’

At about the same time Lidicker’s report was published, Dr. Jack Mahoney at the University of Sydney was studying endemic New Guinea murines with the intention of revising several groups, Pogonomelomys among them. During Musser’s visit to Australia in 1976, Mahoney described his progress with the revision and showed Musser the external, cranial, and dental characters of ruemmleri that set it apart from the other species in Pogonomelomys , features that in combination were not repeated in any other described New Guinea endemic and were going to be used by him to place ruemmleri in its own genus. Mahoney died before completing his revisionary work.

Other systematists working with New Guinea murines were aware of ruemmleri ’s distinct morphology compared to the other species of Pogonomelomys . Ziegler (1982: 880), for example, citing Menzies (1973: 4), indicated that ruemmleri probably merited generic status. Later, Flannery (1990: 239) commented that ‘‘Studies currently underway suggests that ‘‘ P.’’ ruemmleri is the most derived member of a clade containing ‘‘ P.’’ sevia and ‘‘ M. ’’ albidens .’’ Ultimately, Menzies (1990: 132) revised Pogonomelomys , excluding ruemmleri from it and making it the type-species for the new genus Coccymys (‘‘from the Greek coccyx and mys, ‘cuckoomouse’ with reference to spending its early taxonomic life in a place not its own’’), with this diagnosis:

a combination of very long incisive foramina; large 3rd upper molar; incipient division of 1st and 2nd upper molar lophs into separate cusps; very broad interparietal; long palate with deep longitudinal grooves, tail with overlapping scales subtending 3 hairs each, grey-based ventral fur and mammary formula of 1+256.

Menzies provided a description of certain external, cranial, and dental features of C. ruemmleri and compared it with other murines, particularly species of Paramelomys and Pogonomelomys . Pogonomelomys sevia , the other member of Tate’s (1951) ‘‘ sevia-rümmleri group’’ and Flannery’s (1990) suggested ruemmleri-sevia-albidens clade, was extracted from Pogonomelomys by Menzies (1990: 133) and made the type species of Abeomelomys (‘‘from the latin abeo, to go away from’’).

The characterization of ruemmleri by Tate and Archbold and the diagnosis of Coccymys by Menzies were presented in the context of comparisons with other New Guinea murines, primarily species of Melomys and Pogonomelomys , but neither is diagnostic within Indo-Australian Murinae or the geographically broader Murinae . Tate and Archbold’s description of ruemmleri is actually more satisfactory as a diagnosis for Coccymys than is Menzies’s. We expand on their exposition by providing an emended generic diagnosis, and a detailed description of the type species ruemmleri based primarily upon anatomical attributes of dry study skins, cleaned skulls, and some material preserved in fluid.