AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3052, 19 pp., 9 figures, 1 table December 14, 1992

Sucking Lice (Insecta, Anoplura) from Indigenous : a New of Polyplax from a Montane Shrew , and New Information About Polyplax wallacei and P. eropepli

LANCE A. DURDEN' AND GUY G. MUSSER2

ABSTRACT

Polyplax melasmothrixi, a new species of po- from Eropeplus canus from tropical upper mon- lyplacid sucking louse, is described from Melas- tane rain forest also in Central Sulawesi. Host and mothrix naso, a small-bodied shrew rat known habitat associations for these three species ofsuck- only from tropical upper montane rain forest in ing lice are discussed. Polyplax melasmothrixi and Central Sulawesi, Indonesia. The male ofPolyplax P. eropepli are both known only from montane wallacei is described from specimens collected from habitats in Central Sulawesi and both appear to chrysocomus trapped in tropical lowland be host specific (to M. naso and E. canus, respec- evergreen rain forest in Central Sulawesi. A further tively). Contrastingly, P. wallacei parasitizes two specimen ofPolyplax eropepli, a taxon previously species ofBunomys in lowland forests and is known known only from the type series, is documented from North and Central Sulawesi.

INTRODUCTION Melasmothrix naso, Bunomys chrysoco- Musser and Holden, 1991). The shrew rat, , and Eropeplus canus are three murine M. naso, and the large-bodied E. canus have rodents found only in forests on the Indo- been recorded only from montane rainforest nesian island of Sulawesi (Musser, 1987; formations in the mountainous central part

I Assistant Professor and Assistant Curator, Institute of Arthropodology and Parasitology, Georgia Southern Uni- versity, Landrum Box 8056, Statesboro, Georgia 30460. Research Associate, Department ofEntomology, Smithsonian Institution, Washington, DC 20560. 2 Archbold Curator, Department of Mammalogy, American Museum of Natural History.

Copyright C© American Museum of Natural History 1992 ISSN 0003-0082 / Price $2.60 2 AMERICAN MUSEUM NOVITATES NO. 3052 of the island. Bunomys chrysocomus also oc- Information derived from these descrip- curs in montane forest but is most frequently tive surveys will also be significant in a wider encountered at lower altitudes in lowland context. Eventually, phylogenetic data will be tropical evergreen rain forest. These three available for both hosts and parasites, and species of are phylogenetically distant hypotheses focusing on host-parasite coevo- from each other, but they share, in addition lution can be formulated and tested. The re- to their endemicity to Sulawesi, an important lationships between ectoparasites and their trait: they are all parasitized by species of hosts have always intrigued biologists. Re- sucking lice (Anoplura) belonging to the ge- cently, a growing body of literature has de- nus Polyplax. scribed coevolution between groups of hosts The descriptions of these lice, along with and parasites based on comparisons of their notes about the hosts and their hab- phylogenies (Brooks, 1988; Hafner and Nad- itats, form the body ofthe present report. We ler, 1988, 1990; Hellenthal and Price, 1991; describe and illustrate a new species of Po- May and Anderson, 1983; Page, 1991; Toft lyplax based on specimens removed from the and Karter, 1990) as well as apparent coevo- fur of Melasmothrix naso. We also describe lution that in reality reflects host-induced the male of Polyplax wallacei, a louse origi- morphology of the parasite (Downes, 1990). nally known only by female examples (Dur- Preceding this more complex level ofanalysis den, 1987); this ectoparasite has been col- are surveys simply describing the parasites lected from Bunomys chrysocomus trapped and documenting their host species; the re- in lowland evergreen rain forest in north- sults that we present here reflect that level of eastern and Central Sulawesi, and from Bun- inquiry. omys fratrorum, which occurs only in the northeast arm of the island. Finally, we doc- ABBREVIATIONS AND PROCEDURES ument an additional specimen of Polyplax Specimens examined and referred to here eropepli, which until now was only known are deposited in the collections of the Amer- from the type series; this louse is apparently ican Museum of Natural History, New York a specific parasite of Eropeplus canus. (AMNH); the Natural History Museum, Durden and Musser (1991) have summa- London (BMNH); Lance A. Durden (LAD); rized the taxa of sucking lice and their hosts Museum Zoologicum Bogoriense, Bogor, In- so far recorded from Sulawesi. The present donesia (MZB); South Australian Museum, report extends our knowledge beyond that Adelaide, Australia (SAM); and the National summary and is one of a series intended to Museum ofNatural History, Smithsonian In- document the diversity ofectoparasites, some stitution, Washington, D.C. (USNM). ofwhich possess unusual or unique morpho- Other abbreviations used in the descriptive logical traits, found on the array of murine portions of this work are as follows: host species that are endemic to the forests of Sulawesi. This and future papers will con- DAcHS Dorsal accessory head setae DAnHS Dorsal anterior head setae tain descriptions ofnew or poorly known spe- DCAS Dorsal central abdominal setae cies of ectoparasites that have recently been DMHS Dorsal marginal head setae removed from the skins (both dry and pre- DMsS Dorsal mesothoracic setae served in fluid) ofSulawesian rodents housed DPHS Dorsal principal head setae in major museum collections. DPoCHS Dorsal posterior central head setae More than 40 indigenous species of rats DPrS Dorsal paratergal setae and mice have been recorded from Sulawesi DPTS Dorsal principal thoracic setae and its offshore islands (Musser and Holden, DPtS Dorsal prothoracic setae 1991). It is important to document the di- OrS Oral setae versity of ectoparasites and their host asso- SHS Sutural head setae ciations. Results will form another set ofdata SpAtHS Supraantennal head setae StAS Sternal abdominal setae that may be significant for testing hypotheses TeAS Tergal abdominal setae ofzoogeographical relationships among hosts VPaHS Ventral preantennal head setae and their ectoparasites (Durden and Traub, VPHS Ventral principal head setae 1990). VPrS Ventral paratergal setae 1 992 DURDEN AND MUSSER: SUCKING LICE-SULAWESI RODENTS 3

Descriptive format and terminology follow G. Musser in March and April, 1975. Allo- Kim (1966), Kim and Ludwig (1978), and type female and one paratype female from Durden and Musser (1991). Drawings of en- the same host individual as holotype; three tire lice conventionally illustrate dorsal mor- paratype females ex male M. naso (AMNH phology to the left ofthe midline and ventral 225091) at 2255 m; one paratype male ex features to the right. Measurements were male M. naso (AMNH 225105) at 2285 m; made using a calibrated eyepiece micrometer one male ex female M. naso (AMNH 225106) inserted into a high-power phase contrast mi- at 2285 m. In addition, five immature lice croscope. Values of measurements and were removed from AMNH 225106. Allo- weights are reported in millimeters (mm) and type female: MZB. Paratypes: BMNH, LAD, grams (g). All rodent hosts discussed here USNM. (Melasmothrix naso, Eropeplus canus, Bun- DISTRIBUTION: Known by four collections omys chrysocomus, and Bunomysfratrorum) from Melasmothrix naso collected at 2225 are members ofthe subfamily , which and 2285 m (tropical upper montane rain Carleton and Musser (1984) place in the fam- forest) on or close to the summit of Gunung ily of Rodentia. Nokilalaki (2285 m), Central Sulawesi (Su- lawesi Tengah), Indonesia. ACKNOWLEDGMENTS ETYMOLOGY: The species is named for the type host . We are grateful to Michael D. Carleton DIAGNOSIS: Polyplax melasmothrixi is dis- (USNM) and Paula D. Jenkins (BMNH) who tinctive. It is easily distinguished from all arranged for various murine study skins un- other of der their curation to be examined for ecto- species Polyplax by the unusual ap- parasites. Lance Durden's fieldwork in Su- pearance of the male genitalia and by the lawesi was funded by a grant from the setation of gonopods VIII and IX in the fe- Committee for Research and Exploration of male. The following combination of char- the National Geographic Society; he was acters also serves to distinguish this species: sponsored in Indonesia by the Lembaga Ilmu (1) DPHS, DAcHS, and DPoCHS all stout; Pengetahuan Indonesia (LIPI) and the Royal DPHS and DAcHS displaced posterolater- Entomological Society ofLondon. Guy Mus- ally; (2) the shape ofthe thoracic sternal plate; ser's field collections in Sulawesi were partly (3) the presence of some short stout setae, in supported by the Celebes Fund ofthe Amer- addition to longer stout setae, on the abdom- ican Museum as well as Archbold Expedi- inal tergites and sternites; (4) the shape ofthe tions, Inc.; his sponsorship in Indonesia came paratergal plates and the lengths of the setae from LIPI and MZB. He was also assisted by borne on these plates. members ofUnited States Navy Medical Re- DEScRIPTION: Male (fig. 1). Length of ho- search Unit No. 2 (NAMRU-2) in Jakarta. lotype 1.02 mm (mean for series 1.00; range Donna Moore Smith skillfully prepared the 0.96-1.03; N = 3). Head, thorax, and ab- drawings included in this paper. Drs. Richard domen well sclerotized. G. Robbins and Nixon Wilson critically re- Head. About as wide as long with blunt viewed an earlier draft of the manuscript. anterior apex; 2 DAnHS, 1 DMHS, 1 OrS, 1 SHS (2 present on 1 side of 1 specimen), 1 Polyplax melasmothrixi, new species SpAtHS, and 2 VPaHS on each side; DPHS relatively short and stout, extending about HOLOTYPE: Male collected by L. A. Durden halfdistance to thoracic spiracle, with 1 stout from pelt of adult male Melasmothrix naso DAcHS distal to DPHS on each side; both (AMNH 225100) collected by G. G. Musser DPHS and DAcHS displaced posteromargin- at 2255 m on Gunung Nokilalaki (1°16'S, ally; 1 stout peglike DPoCHS on each side; 12001 0'E), Central Sulawesi, Indonesia on 29 VPHS long and fairly stout. Antennae 5-seg- April, 1975. Deposited in the MZB. mented with basal segment much larger than REFERRED SPECIMENS: Seven additional second segment, wider than long; third an- adult lice (2 males, 5 females) all removed tennal segment with anterior projection as by L. A. Durden from pelts of M. naso from typical for males of Polyplax spp. trapped on Gunung Nokilalaki by G. Thorax. Broad with almost parallel outer 4 AMERICAN MUSEUM NOVITATES NO. 3052

and hind coxae subtriangular; forelegs small, each with narrow acuminate claw; mid and hind legs progressively larger and with cor- respondingly larger acuminate claws. Abdomen. Slightly wider than thorax; one broad plate per segment dorsally (on seg- B ments 2-8) and ventrally (on segments 4-7); two narrow plates ventrally on each of seg- ments 2 and 3; curving, concave plate ven- trally on segment 8; most plates with finely crenulated posterior margins; dorsal plates on segments 2-7 each with 6-8 thickened TeAS; dorsal plate on segment 8 with 2 long TeAS; ventral plates 1-3 and 8 each with 3- 4 StAS; ventral plates 4-7 each with 6-7 thickened StAS; ventral plate 9 with 2 long marginal StAS; some TeAS and StAS short and stout. Paratergal plates (fig. 1C) present on segments 2-8; all plates with 2 apical setae each, with DPrS slightly longer than corre- sponding VPrS of same plate: plates I and II with short setae, plates III-V with setae of intermediate length, and plates VI and VII with longer setae; all plates subtriangular; plates IV and V with both apical angles pro- duced into points; plates II-VII each with moderately sized spiracle. Genitalia (fig. ID). Basal apodeme deeply concave posteriorly and with differential scle- rotization; parameres flanged medially, notched anteriorly, and appearing to articu- late with posterior projections of basal apo- deme; pseudopenis short, extending approx- imately to apices ofparameres, and with three sclerotized patches. Female (fig. 2). Length ofallotype 1.18 mm (mean for series 1.18; range 1.14-1.24; N = 5). D Head, thorax, and legs. As in male unless indicated otherwise. Third antennal segment not modified; thoracic sternal plate (fig. 2B) posteriorly more prolonged than in male. Fig. 1. Polyplax melasmothrixi, male. A, hab- Abdomen. With one broad plate dorsally itus; B, thoracic sternal plate; C, paratergal plates; on each of segments 2, 3, and 8, and two D, genitalia. broad plates dorsally on each of segments 4- 7; two broad plates ventrally on each of seg- margins; thoracic sternal plate (fig. 1B) ta- ments 2-7; dorsal plates 1, 10, and 11 each pering to slightly prolonged, rounded, pos- with four thickened TeAS; dorsal plates 2-9 terior apex, with central, blunt, anterior pro- each with 6-7 thickened TeAS; ventral plate jection, and slightly concave, lateral margins; 1 with two thickened StAS, ventral plate 2 mesothoracic spiracle moderate in size (0.018 with four thickened StAS, and ventral plates mm in diameter); DPTS short (0.098 mm); 3-12 each with 5-7 thickened StAS; StAS 1 DMsS and 1 DPtS on each side. Legs. Mid elongated on ventral plate 12; some TeAS 1 992 DURDEN AND MUSSER: SUCKING LICE-SULAWESI RODENTS 5 and StAS short and stout; most tergal and sternal abdominal plates with fine crenula- tions along posterior margins. Paratergal plates (fig. 2C) present on segments 2-8 and all bearing 2 apical setae each; DPrS longer than corresponding VPrS on plates I, II, IV, and V; plates I-III with short apical setae, plates IV and V with setae of intermediate length, and plates VI and VII with longer B setae; plates I and II subtriangular; plates II- VI with both apical angles produced into points; plates II-VII each with moderately sized spiracle. Genitalia (fig. 2D). Subgenital plate sub- triangular; gonopods VIII each with one long lateral seta and one short seta situated close to center ofgonopod; gonopods IX each with one short stout seta but with smaller anterior lobe also bearing short seta; vulvar fimbriae distinct. REMARKS: Morphologically, P. melas- mothrixi is an unusual louse when compared to its congeners. Some head setae and most abdominal setae are stout, particularly in the male. Both Polyplax phthisica Ferris and P. smallwoodae Johnson, which are parasites of African murines (Lemniscomys spp.), simi- larly possess stout abdominal setae (Johnson, 1960) but otherwise these lice do not appear to be closely related to P. melasmothrixi. Genitalic characters ofP. melasmothrixi also are unusual. The male pseudopenis is unique in possessing three sclerotized distal patches. C Similarly, the proximally notched parameres that appear to articulate with the distal arms of the basal apodeme are very unusual. The setation of the female gonopods VIII and IX is also distinct. These and other characters strongly suggest that P. melasmothrixi is morphologically unique and not closely related to any other known species in the genus Polyplax. The host species, M. naso, is unique from two Fig. 2. Polyplax melasmothrixi, female. A, perspectives. It is the only known member habitus; B, thoracic sternal plate; C, paratergal ofMelasmothrix. Its closest phylogenetic rel- plates; D, genitalia. atives are the small-bodied shrew rats Ta- teomys rhinogradoides and T. macrocercus, also endemic to the mountains ofCentral Su- lawesi, and they along with M. naso form a pect the morphology of sucking lice from the monophyletic group without apparent close other two species of shrew rats to resemble relatives either on Sulawesi or elsewhere in that characterizing Polyplax melasmothrixi; the Indo-Australian region (Musser, 1982; unfortunately, no lice were found on any of Breed and Musser, 1991). We would also ex- the preserved specimens of . 6 AMERICAN MUSEUM NOVITATES NO. 3052

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Fig. 4. Melasmothrix naso, the host of Polyplax melasmothrixi. An adult male caught during the day on the summit of Gunung Nokilalaki in habitat similar to that shown in figure 3.

HOST HABITAT: All three species of Su- Melasmothrix naso is small in body size lawesian small-bodied shrew rats are known with an elongate head and muzzle, small eyes to occur only in the mountainous central core and ears, chunky body, short tail, elongate of Sulawesi. Records of Melasmothrix naso claws on the front feet, and velvety dark are from 6000 ft at Rano Rano (1°30'S, chestnut fur (fig. 4). It is diurnal and terres- 120°28'E) where the holotype (USNM trial, uses mossy and earthen runways con- 219752) was obtained (Miller and Hollister, cealed beneath moss-covered rotting treefalls 1921), and between 6400 and 7500 ft on the or root entanglements, and apparently feeds southwest slope and near the summit ofGun- primarily on earthworms and fungus gnats ung Nokilalaki (1°16'S, 120°10'E) where 35 (Musser, 1982). specimens were collected (Musser, 1982). Cool, wet, and mossy forest above 6000 ft Polyplax wallacei Durden is the general habitat where all examples of Polyplax wallacei Durden, 1987: 812. M. naso have been encountered (fig. 3). Polyplax wallacei Durden. Durden and Traub, Among the characteristics of this formation 1990: 57 (zoogeography). are the short canopy; small trees representing Polyplax wallacei Durden. Durden and Musser, species associations and morphologies dis- 1991: 7 (faunal relationships). tinctive to mountain environments; low di- HOLOTYPE: Female designated by Durden versity in species oftrees, shrubs, palms, and (1987). In USNM. Allotype male removed other plants; dense moss cover; and high fre- by L. A. Durden from pelt ofa male Bunomys quency and long duration of rain and mist. chrysocomus (AMNH 226924) collected by Quantitative details, other descriptive obser- G. G. Musser at 1000 m near Tomado (1° 9'S, vations, and other places where the shrew rats 120°03'E), Danau (Lake) Lindu, Central Su- were trapped are documented by Musser lawesi, Indonesia on 22 July, 1973. Depos- (1982: 71). ited in the MZB. 8 AMERICAN MUSEUM NOVITATES NO. 3052

B

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D Fig. 5. Polyp/ax wallacei, male. A, habitus; B, thoracic sternal plate; C, paratergal plates; D, genitalia.

REFERRED SPECIMENS: Eight additional lice bearing the same locality data as the allotype (4 males, 4 females). Of these, an additional host but collected on 21 July, 1975. The re- male and one female were removed from the maining five specimens, most of which are same pelt from which the allotype was taken. in relatively poor condition, were located in One male louse was removed from the pelt the USNM slide-mounted Anoplura collec- ofa B. chrysocomus female (AMNH 223038) tions where they had been misidentified as 1 992 DURDEN AND MUSSER: SUCKING LICE-SULAWESI RODENTS 9

Polyplax spinulosa Burmeister. These speci- Thorax. Slightly broader than long; tho- mens were also collected from Tomado; one racic sternal plate (fig. SB) distinctive with male and two female lice were collected on rounded, central, anterior apex, angulate less 22 January, 1972 and bear the field collection well-developed anterolateral angles, and ta- number DJ.M-2366, while two male lice were pering to broadly rounded posterior apex; collected on 24 January, 1972 (field collec- mesothoracic spiracle small (0.016 mm in tion no. DJ.M-2455). Hosts for both ofthese diameter); DPTS moderate in length (0.115 collections were originally labeled on the mm), extending beyond insertions of first slides as " penitus penitus," which we DCAS; Legs. With subtriangular coxae; fore- have since determined to be Bunomys chry- legs with small acuminate claw; hindlegs socomus. Both ofthese 1972 collections were slightly larger than midlegs, both with strong, made by members of NAMRU-2 teams. pointed claws. Paratypes: BMNH, LAD, MZB, USNM. Abdomen. Wider than thorax; two anterior DISTRIBUTION: Known from both North rows of 2 DCAS followed by one broad plate and Central Sulawesi (Sulawesi Utara and Su- dorsally on each ofsegments 2-8; dorsal plates lawesi Tengah, respectively) from two species 1-6 (on abdominal segments 2-7) each with offorest rats belonging to the genus Bunomys. 8-10 TeAS; dorsal plate 7 (on abdominal seg- All known collections from North Sulawesi ment 8) with 4 TeAS; two narrow plates ven- are from Dumoga-Bone National Park trally on each of segments 2 and 3, and one (0034'N, 123054'E), at an altitude of 230 m, broad plate ventrally on each of segments 4- while those from Central Sulawesi all origi- 7; ventral plates 1 and 3-7 each with 5-7 nate from Tomado near Danau (Lake) Lindu StAS; ventral plates 2 and 8 each with 4 StAS; (elevation 1000 m). some TeAS and StAS short. Paratergalplates DIGANosIs: The female of P. wallacei was (fig. 5C) present on segments 2-8; plates I-V described and diagnosed by Durden (1987). subtriangular, each with two short apical se- The male, described in this paper, has mor- tae; plates VI and VII each with two long phological similarities to Polyplax spinulosa apical setae; DPrS longer than corresponding Burmeister, a widespread parasite of domes- VPrS on plates I-III; plates I-VI with dorsal tic Rattus. The male of P. wallacei can be angles developed into points; plates II-VII distinguished from P. spinulosa and from each with moderately sized spiracle. other species of Polyplax by a combination Genitalia (fig. 5D). Subgenital plate with ofthe following traits: (1) distinctively shield- medial lacuna and one long lateral seta on shaped thoracic sternal plate with extended each side of plate immediately anterior to anterolateral angles (fig. SB); (2) the shapes lacuna; basal apodeme much longer than par- and lengths of the attached apical setae for ameres, forked posteriorly; parameres the paratergal plates (fig. SC); (3) the shapes smoothly curved; pseudopenis thick, extend- and proportions ofthe genitalic structures (fig. ing appreciably beyond apices of parameres. SD). REMARKS: In addition to the new records DEscRIPrIoN: Male (fig. 5). Length of al- from Central Sulawesi recorded here, Durden lotype 0.98 mm (mean for series 0.98; range (1987) collected the following specimens of 0.97-1.00; N = 5). Head, thorax, and ab- P. wallacei from tropical lowland evergreen domen well sclerotized. rain forest (altitude 230 m) in Dumoga-Bone Head. About as wide as long with anterior National Park, North Sulawesi, during Feb- apex broadly rounded; 2 DAcHS (both an- ruary 1985: teromedial to DPHS), 2 DAnHS, 2 DMHS, 1 DPoCHS, 3 OrS, 3 SHS, and 2 SpAtHS on each side; DPHS almost extending to tho- 2 females ex male B. chrysocomus (LAD no. 24), racic spiracle; VPHS short and close to in- 2 females ex male B. chrysocomus (LAD no. 32), sertion of first antennal segment. Antennae 2 females ex female B. chrysocomus (LAD no. 51), 5-segmented 4 females ex female B. chrysocomus (LAD no. 69), with first segment much larger 1 female ex female Bunomysfratrorum (LAD no. than second, about as wide as long; third seg- 43), ment prolonged anterolaterally as is charac- 1 female ex female B. fratrorum (LAD no. 53), teristic of male Polyplax. 1 female ex female sp. (LAD no. 73). 10 AMERICAN MUSEUM NOVITATES NO. 3052

Unfortunately, the last host specimen listed Additional survey of the other species of (LAD no. 73) was not retained as a voucher Bunomys is needed to determine what kinds but our subsequent examination ofcollection ofsucking lice they may harbor. For example, records indicates this was not a spe- in Central Sulawesi, B. andrewsi occurs in cies of Taeromys. Consequently, all reliable tropical lowland evergreen rain forest, and B. known host records for P. wallacei pertain to penitus is the common Bunomys in montane Bunomys chrysocomus and B. fratrorum. habitats except where the mountain B. pro- Polyplax wallacei may be restricted to pop- latus is found (Musser, 1991; Musser and ulations of Bunomys living at low altitudes, Holden, 1991). Samples of B. chrysocomus at least between 230 and 1000 m, the ele- have been taken at some of the same places vations from which the louse has been re- that yielded specimens of B. penitus and B. corded. The known hosts, however, are not andrewsi (Musser, MS), and close to B. pro- similarly ecologically bracketed. In Central latus (Musser, 1991). Is Polyplax part oftheir Sulawesi, samples of Bunomys chrysocomus ectoparasite fauna and, ifso, do they all share were collected by Musser from 300 m up the same species ofsucking louse? IfPolyplax through lowland evergreen rainforest to the wallacei is actually restricted to hosts living interface with tropical lower montane rain at low and middle altitudes, we would not forest (about 1300 m) and higher into mon- expect to find it parasitizing either B. penitus tane habitats between 1370 and 1515 m or B. prolatus but would not be surprised if (AMNH 223566-223568, 223570, 225148 it occurred on B. andrewsi, which is mor- are the montane voucher specimens). The phologically more similar to B.fratrorum than species has also been obtained in montane to any other species of Bunomys (Musser, formations up to 6000 ft on Gunung Tam- MS). bussissi (Musser, 1991), above 6000 ft on Another facet of the association between Gunung Lehio (USNM 218673 and 218681), P. wallacei and Bunomys is the low frequency at 2200 m on Pegunungan Latimojong of louse infestation on B. fratrorum. Durden (AMNH 196578-196584) in the central part (1987: 814) noted that B. chrysocomus ap- of the island, and at 1500 and 2000 m at peared to be the primary host of P. wallacei Tanke Salokko on Pegunungan Mekongga in and that its occurrence on B.fratrorum "could the southeastern arm (Musser, 1991). Dur- represent accidental infestations through eco- den unsuccessfully sought lice from the fur logical associations." This suggestion could of these montane samples, which consisted be tested by surveying freshly caught B. fra- ofstudy skins and fluid-preserved specimens. trorum specimens from localities where it does In contrast to Bunomys chrysocomus, which not occur together with B. chrysocomus. Also, occurs throughout most of Sulawesi (at least if B. chrysocomus is really the primary host where collectors have sampled) between about and accidental infestation of other species is 300 and 1800 m (Musser, MS), B. fratrorum a possibility, P. wallacei might also be found has been found only in the northeastern arm on other species of Bunomys that are sym- ofSulawesi east ofthe Gorontalo region. Most patric with B. chrysocomus. samples come from lowlands and middle al- Whether or not B. fratrorum is an acci- titudes, but the species also reaches montane dental or primary host ofP. wallacei, because forest environments as documented by sam- the louse has been found on that host and ples from 1780 m on Gunung Maujat (SAM not on a murine in another genus suggests m12644 and m12681) and 2000 m on the evolutionary specificity between P. wallacei slopes of Gunung Klabat (USNM 217581, and the genus Bunomys. Throughout its geo- 217582, and 217585). We have not surveyed graphic range, B. chrysocomus occurs in the these animals for lice. That Polyplax wallacei same habitat and in Central Sulawesi was col- may be restricted to habitats at low and mid- lected at the same sites as Rattus hoffmanni, dle altitudes in tropical lowland evergreen Maxomys hellwaldii, and M. muschenbroe- rain forest while their hosts are not so con- kii, other Sulawesian endemic murines, but fined is the obvious hypothesis to be tested those hosts are parasitized by species of Ho- by sampling freshly caught rats collected in plopleura, not Polyplax (Durden, 1990; Dur- montane situations. den and Musser, 1991). 1 992 DURDEN AND MUSSER: SUCKING LICE-SULAWESI RODENTS I1I

Fig. 6. Bunomys chrysocomus, the host ofPolyplax wallacei.. An adult female from Sungai Tokararu, 1150 m.

HOSTS AND HABITAT: Bunomys chrysoco- In its morphology and external appear- mus is of medium body size (see measure- ance, B. fratrorum is a larger version of B. ments listed in tables 1 and 2 in Musser, chrysocomus (compare the values listed in 1991), with a tail shorter than head and body, tables 1-3 in Musser, 1991), and examples small eyes and ears, a long snout, and covered of it have often been misidentified as the by dark, soft fur (fig. 6). Musser (fieldnotes smaller species in museum collections. In ad- in AMNH) found it to be an opportunistic dition to its much larger size, B. fratrorum feeder, subsisting on insects, earthworms, has a longer tail relative to head and body snails, small frogs and lizards, and sometimes length, the distal portion of the tail is white fruit. in most specimens, and cusp patterns ofmo- In Central Sulawesi, Musser encountered lars are less complex. Bunomys fratrorum B. chrysocomus most commonly in the wetter likely occurs in similar habitat as B. chryso- and cooler parts oftropical lowland evergreen comus because specimens of both have been rain forest (fig. 7), usually under dense cover collected at the same place (although no in- along the sides of wet ravines and small formation on microhabitats was provided), streams. It was rarely collected higher on and they may have a similar diet. The stom- slopes, ridgetops, or beneath open understo- ach of one specimen we studied contained ry, where the ambient environment is slightly remains of insects and earthworms. drier than that characteristic of ravines and streamside habitats. At higher elevations within the boundaries oftropical lower mon- Polyplax eropepli (Ewing) tane rain forest, B. chrysocomus was also Eremophthirius eropepli Ewing, 1935: 209. found in wetter areas of the forest with good Polyplax eriopepli [sic] Ewing. Ferris, 1951: 207 cover. (catalog). 12 AMERICAN MUSEUM NOVITATES NO. 3052

Fig. 7. Lowland evergreen rain forest along Sungai Tokararu at 1150 m, Central Sulawesi, 1973. Examples of Bunomys chrysocomus were trapped beneath the dense streamside vegetation. 1 992 DURDEN AND MUSSER: SUCKING LICE-SULAWESI RODENTS 13

Polyplax eropepli (Ewing). Johnson, 1958a: 47 1981) as well as spermatozoal morphology (lectotype designated). (Breed and Musser, 1991). Lenomys is rep- Polyplax eropepli (Ewing). Durden, 1087: 814 (re- resented by few specimens in museums, and description). unfortunately we have not found lice on any Polyplax eropepli (Ewing). Durden aind Traub, of those we examined. 1990: 57 (zoogeography). HOST AND HABITAT: Eropeplus canus is a Polyplax eropepli (Ewing). Durden and Musser, member of the montane murine fauna en- 1991: 7 (faunal relationships). demic to Central Sulawesi (Musser and Hold- REFERRED SPECIMEN: One female louse re- en, 1991: 406). Although infrequently en- moved by L. A. Durden from the pelt of an countered by collectors, it is documented by Eropeplus canus female (USNM 219711) 13 examples from the following places: Gun- trapped at 1800 m at Rano Rano, Central ung Nokilalaki (1°16'S, 120°10'E) at 5950 ft Sulawesi, Indonesia, by H. C. Raven on 15 (AMNH 225119 and 225120), 7200 ft December, 1917 (Miller and Hollister, 1921). (AMNH 223553, 223554, 223557), 7250 ft Deposited in the USNM. (AMNH 225121-225123), and 7500 ft REMARKS: Until now, P. eropepli was (AMNH 223555, 223556, 225124); Rano known only from four specimens (two males, Rano (1°30'S, 120°28'E) at 6000 ft (USNM two females) removed from the same pelt 219711); Gunung Lehio (1°33'S, 110°53'E) listed here for the new specimen (Ewing, 1935; above 6000 ft (USNM 218707, holotype); Ferris, 1951; Johnson, 1958a; Durden, 1987). Pegunungan Quarles, Bulu Karua (2°56'S, These four specimens are mounted on the 119039'E) at 6000 ft (BM 40.386 and 40.387); same microscope slide, which bears the and Pegunungan Latimojong (3°30'S, USNM type number 44906. All five P. ero- 121023'E) at 2200 m (AMNH 196592). pepli specimens are in poor condition, but Eropeplus canus is large and bulky with a the new specimen has retained some intact very long bicolored tail (fig. 8). Among five structures that are damaged in the type series. adults from Gunung Nokilalaki, the range in Both female paralectotypes are missing all leg length of head and body is 230-255 mm, segments other than the coxae; the new spec- length of tail is 270-296 mm, length of hind imen, however, has intact legs and these do foot is 46-48 mm, length ofear is 21-27 mm, not differ significantly from those illustrated and weight is 190-315 g. Long and thick gray by Durden (1987) for the male. Other differ- fur covers the body and proximal portion of ences noted for the new specimen are as fol- the tail. lows: (1) a small, anterior, central projection Tropical montane rain forest (generally de- is present on the thoracic sternal plate; (2) on scribed by Whitmore, 1984) is the general paratergal plate VII, one apical seta remains habitat ofE. canus (fig. 9). Musser (1982) and intact; it is much longer than the correspond- Musser and Dagosto (1987) provide descrip- ing setae on the paralectotypes suggesting se- tions of this environment on Gunung Nok- tal damage in those specimens; (3) on gon- ilalaki. Musser trapped the rats on wet ground opods VIII, the lateral setae (broken on both near and within rock clusters covered with sides in the paralectotypes) are long, extend- moss and held together by tree roots, along- ing beyond the outline of the abdomen for side rotting trunks, and 15 ft above ground about 40% of their length; (4) on each gon- on a woody vine. Fruits, leaves, palm hearts, opod IX, a single stout seta, rather than two and some kinds ofinsects were eaten by cap- narrow setae, is present. tive rats (Musser, fieldnotes in AMNH). Polyplax eropepli has never been collected from murine species other than Eropeplus ASSOCIATED MORPHOLOGICAL canus and it is probably host specific to that DIVERGENCE IN animal. However, the uniqueness of this as- sociation has to be tested by surveying Len- HOST AND PARASITE omys meyeri for sucking lice. That murine is Because evolutionary parallelism at the endemic to Sulawesi and is the closest known levels of genera and species is a relationship relative of E. canus, as revealed by shared between sucking lice that is widespread (Kim, distinctive cranial and pelage traits (Musser, 1985, 1988), we might expect each species of 14 AMERICAN MUSEUM NOVITATES NO. 3052

Fig. 8. Eropeplus canus, the host of Polyplax eropepli. An adult female from near the summit of Gunung Nokilalaki, 7500 ft. louse associated with the species of Melas- P. spinulosa, a louse found on commensal mothrix, Eropeplus, and Bunomys to also be Rattus species (table 1; Durden, 1987; Dur- unique, possibly at the generic level. Melas- den and Page, 1991) that occur on Sulawesi mothrix naso, the louse found on the shrew but are not native to the island (Musser and rat, M. naso, is diagnosed by a set of char- Holden, 199 1). Polyplax wallacei, which par- acters unique to it (stout setae, male pseu- asitizes Bunomys chrysocomus and B. fratro- dopenis with three sclerotized distal patches, rum, also resembles P. spinulosa in some fea- proximally notched parameres that articulate tures and shares one trait (the general shape with distal arms of the basal apodeme, and ofthe thoracic sternal plate) with P. cutchicus, characteristic setation of female gonopods a parasite ofthe Indian rat cutch- VIII and IX), suggesting remote affinity with icus (table 1; Durden, 1987). any other described relative. However dis- These relationships suggest that the re- tinct it is, this louse does not appear to be ported evolutionary parallelism between part of a monophyletic group different from sucking lice and host taxa cited by Kim (1988) that formed by other species of Polyplax, a does not describe the discordant degree of new monotypic genus, for example. morphological divergence that exists be- Polyplax eropepli, the sucking louse spe- tween host and louse among certain species cific to Eropeplus canus, is not as distinctive of Indo-Australian murine rodents. Melas- in its morphology as is P. melasmothrixi. mothrix naso is in a clade with two other Some characters ofP. eropepli are shared with species of small-bodied shrew rats endemic 1992 DURDEN AND MUSSER: SUCKING LICE-SULAWESI RODENTS 15

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.0 TABLE 1 Principal Hosts and Known Geographical Distributions ofPolyplax spp. Sucking Lice Parasitic on Murine Rodentsa Louse species Host species Distribution Reference P. abyssinica Arvicanthus niloticus N. & E. Africa Johnson (1960) P. acomydis Acomys spinosissimus Mozambique Kim and Emerson (1970) P. antennata Apodemus mystacinus E. Europe Smetana (1960) P. arvicanthis Rhabdomys pumilio E. & Southern Africa Johnson (1960) P. asiatica indica Southern Asia & Near East Durden et al. (1990) P. blanfordi Cremnomys blanfordi Mishra (1981) P. brachyrrhyncha Acomys cahirinus N. Africa to Pakistan Durden (1991) P. bullimae bagopus Philippines: Mindanao Johnson (1958b) P. cummingsi Dasymys spp. Africa Durden (1991) P. cutchicus Cremnornys cutchicus India Mishra (1981) P. dacnomydi Dacnomys millardi China: Yunnan Chin (1990) P. dolichura Acomys cahirinus Sudan Johnson (1962b) P. eropepli Eropeplus canus Indonesia: Sulawesi Durden and Musser (1991) P. expressa "Rattus sp." Philippines: Luzon Johnson (1964) P. gracilis minutus Eurasia Beaucournu (1968) P. grammomydis dolichurus Southern Africa Johnson (1960) P. hoogstraali Acomys spp. Egypt Johnson (1960) P. humae Cremnomys blanfordib Pakistan Khan and Khan (1985) P. indica Golunda ellioti India Mishra (1981) P. insulsa Leopoldomys sabanus Malaysia, Natuna Is. Johnson (1964) L. edwardsic China Chin (1980) P. kondana kondana India Mishra (1981) Millardia meftada Pakistan Durden et al. (1990) P. melasmothrixi Melosmothrix naso Indonesia: Sulawesi Present report P. meridionalis Acomys spinosissimusd Botswana Johnson (1962a) P. oxyrrhyncha Acomys spp. N. & E. Africa Durden (1991) P. phloeomydis Phloeomys cumingi Philippines: Luzon Cuy (1982) P. phthisica Lophuromys spp. E. & Central Africa Durden (1991) P. praomydis Aethomys namaquensis S. Africa & Namibia Johnson (1960) P. pricei Niviventer confucianuse N. Thailand Kim (1968) P. serrata Mus musculus Cosmopolitan Durden et al. (1990) P. smallwoodae Lophuromys spp. W. & Central Africa Durden (1991) P. solivaga Aethomys chrysophilus S. Africa Johnson (1962b) P. spinulosa Rattus norvegicus, Cosmopolitan Durden (1991) R. rattus Rattus argentiventer S.E. Asia to New Guinea Durden (1987) Rattus exulans Indo-Pacific region Durden (1987) P. tarsomydis apoensis Philippines: Mindanao Ewing (1935) P. thamnomydis Grammomys rutilans Central Africa Johnson (1960) P. vicina Rattus losea f China Blagoveshchensky (1972) P. visenda Leopoldomys sabanus f Vietnam Blagoveshchensky (1972) P. wallacei Bunomys chrysocomus Indonesia: Sulawesi Durden (1987) P. waterstoni Mastomys spp., Southern Africa Durden (1991) Praomys spp. a An additional 39 species of Polyplax are known. Collectively, they parasitize belonging to eight other subfamilies of Muridae (in the sense of Carleton and Musser, 1984) and to the families Sciuridae and Soricidae. b There are no substantiated records of Cremnomys blanfordi from Pakistan. This murine is endemic to peninsular India and to Sri Lanka. We assume that the correct type host of Polyplax humae is either Millardia gleadowi or M. meltada. c We have not seen the louse or host material reported by Chin (1980) for P. insulsa from L. edwardsi. d The type host ofPolyplax meridionalis was listed as Acomys cahirinus by Johnson (1962a). However, A. spinosissi- mus is the only murine of the genus Acomys that is known from Botswana (Dippenaar and Rautenbach, 1986). e The type host of Polyplax pricei was listed as Rattus niveiventer [sic] (=Niviventer niviventer). This murine does not occur in northern Thailand (Marshall, 1977) and we have since determined the type host to be Niviventer confucianus. fWe have not seen the louse or host material reported by Blagoveschensky (1972). 1992 DURDEN AND MUSSER: SUCKING LICE-SULAWESI RODENTS 17 to Sulawesi, Tateomys rhinogradoides and T. Breed, W. G., and G. G. Musser macrocercus. Eropeplus canus and Lenomys 1991. Sulawesi and Philippine rodents (Mur- meyeri form a separate clade that is phylo- idae): a survey of spermatozoal mor- phology and its significance for phylo- genetically distant from the group of shrew genetic inference. Am. Mus. Novitates rats (Musser, 1982; Breed and Musser, 1991). 3003: 15 pp. Bunomys and Rattus are part of a clade that Brooks, D. R. includes genera that are native not only to 1988. Macroevolutionary comparisons of host Sulawesi but to other islands and continents and parasite phylogenies. Annu. Rev. in the Indo-Australian region (Musser and Ecol. Syst. 19: 235-259. Heaney, 1992; Kitchener et al., 1991), and Carleton, M. D., and G. G. Musser one that is not phylogenetically closely relat- 1984. Muroid rodents. In S. Anderson and J. ed to either the shrew rat or Eropeplus-Len- K. Jones (eds.), Orders and families of omys clades. Similarly, the Indian rat, Crem- Recent mammals ofthe world, pp. 289- not related to 379. New York: Wiley. nomys cutchicus, is closely any Chin, T.-H. of the endemic Sulawesi murines. Yet it and 1980. Studies on Chinese Anoplura V. The species in the other three clades are parasit- polyplacid genera Polyplax, Haemodip- ized by species of sucking lice in the same sus and Sathrax. Acta Acad. Med. Gui- supposed monophyletic group, Polyplax. yang 5:101-107. [In Chinese with En- These and other known examples of murine/ glish summary] Polyplax host-parasite associations are listed 1990. Polyplax dacnomydi-a new species of in table 1. Most of the lice shown in table 1 sucking louse (Anoplura: Polyplacidae) are clearly host specific (monoxenous). How- from Yunnan. Entomotaxonomia 12: ever, because diverse murine hosts with var- 5-7. [In Chinese with English summary] ious Old World distributions (except for two Cuy, L. S. 1982. A new species of louse (Phthiraptera: species that have accompanied their cos- Hoplopleuridae), with notes on its host mopolitan hosts around the globe) are par- from the Philippines. Kalikasan, Phil- asitized, host-parasite coevolution in the strict ippine J. Biol. 11: 199-202. sense (i.e., Fahrenholz' rule) does not appear Dippenaar, N. J., and I. L. Rautenbach to have occurred at the genus and species 1986. Morphometrics and karyology of the levels. These observations are based on mor- southern African species of the genus phological characters and need to be com- Acomys (Rodentia, Muridae). Ann. pared with results from analyses ofother kinds Transvaal Mus. 34: 129-183. of systems, molecular traits for example. Downes, B. J. Some other ectoparasites ofSulawesi rodents 1990. Host induced morphology in mites: im- plications for host-parasite coevolution. may show clearer host-parasite morpholog- Syst. Zool. 39: 162-168. ical coevolution. For example, many unde- Durden, L. A. scribed genera and species of Sulawesian ro- 1987. The genus Polyplax (Anoplura: Poly- dent fleas are known, and distinct flea genera placidae) from Sulawesi, Indonesia, with appear to parasitize distinct murine genera the description of a new species. Proc. in many cases (Durden and Traub, 1990; R. Entomol. Soc. Washington 89: 811-817. Traub, personal commun.). 1990. The genus Hoplopleura (Anoplura: Ho- plopleuridae) from murid rodents in Su- lawesi, with descriptions of three new REFERENCES species and notes on host relationships. J. Med. Entomol. 27: 268-281. Beaucournu, J.-C. 1991. New records of sucking lice (Insecta: 1968. Les anopluores de lagomorphes, ron- Anoplura) from African mammals. J. geurs et insectivores dans la region Pa- African Zool. 105: 331-342. learctique Occidentale et en particulier Durden, L. A., and G. G. Musser en France. Ann. Parasitol. Hum. Comp. 1991. A new species ofsucking louse (Insecta, 43: 201-271. 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