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Pacific Science (1991), vol. 45, no. 2: 123-130 © 1991 by University ofHawaii Press. All rights reserved

Origin, Dispersal Routes, and Geographic Distribution of exulans, with Special Reference to !

MERE ROBERTS2

ABSTRACT: Current anthropological theory and archaeological evidence have led to a reappraisal ofthe hypothesized route ofdispersal of the Polynesian , Rattus exulans, through the Pacific and to New Zealand. This commensal seems to have entered the -Tonga-Samoa region by way of , in association with the people of the Lapita cultural complex. The hypothesized migrations ofthose people and hence presumably ofR. exulans throughout Near and Remote are presented here, along with a brief review of this rat's history and current distribution in New Zealand, the last settled and southern­ most landmass in its range.

THE WIDESPREAD AND UNIFORM DISTRIBUTION accounts in the literature (Tate 1935, Schwarz of the , Rattus exulans (Peale), and Schwarz 1967) that deal with the origin throughout the insular Pacific is at variance and dispersal of this both reflect the with the observed distribution of noncom­ then-prevailing anthropological view (e.g., mensal ; this has given rise to the Buck 1938) that the Proto-Polynesians hypothesis that its dispersal occurred in asso­ entered the Pacific by way of and ciation with human movements. The archaeo­ not Melanesia. For example, after studying logical evidence supports this theory; for the comparative anatomy and morphology of instance, the Pleistocene mammalian fauna numerous specimens ofthe Polynesian ratcol­ of includes two fruit bats, a lected from throughout its geographical phalanger, a wallaby, Rattus mordax, and range, Tate (1935) related this species to the Rattus praetor. Rattus exulans appears later, concolor group ofRattus, whose geographical along with the commensal pig, dog, and range includes the Malay states, Sumatra, , at around 3100-2500 Before Present Java, Borneo, the , Celebes, New (B.P.). The temporal specificity ofartifacts and Guinea, and the Pacific islands including New faunal remains found in the sites led Allen et Zealand, but excluding mainland . al. (1989) to conclude that apart from the He went on to suggest that speciation of bats, all appear to be human introductions. R. exulans occurred in the Philippine­ Anthropological information also suggests Borneo-Java region, and from there this rat that the dispersal of R. exulans through­ dispersed eastward into the Pacific (Figure I). out the Pacific took place (accidentally or He pointed out that "since all have been deliberately) in association with the migra­ carried by man, whether voluntarily or in­ tions of the ancestral peoples of this region. voluntarily, they must have followed his col­ From the New Zealand perspective then, onization tracks" and that the "original questions concerning the origin of this rat course from the mainland [of Southeast ] become a question ofthe origin ofthe Maori, was probably from Borneo and the Philip­ and hence of the Polynesian peoples. Based pines via the Caroline Islands rather than on this presumed relationship, two previous through and the Solomon Is­ lands" (Tate 1935: 169). Schwarz and Schwarz (1967) suggested that the origin of the R. exulans series ofcommensals was from 1 Manuscript accepted 2 May 1990. 2 Department of Zoology, University of Auckland, a wild subspecies, Rattus rattus wichmanni, of Auckland, New Zealand. in the (Figure 1). 123 124 PACIFIC SCIENCE, Volume 45, April 1991

20'

• CHRISTMAS I 0'

MARQUESAS

20' AUSTR .... LI ....

/ '. RAP" I '/ i

o 2000k dtt~'~'~o .0'

o 180· '40·

FIGURE 1. Hypothesized insular Pacific dispersal routes of Rattus exulans. Solid lines modified after Tate (1935); dashed lines according to Schwarz and Schwarz (1967).

Divergence and spread then occurred to the have occurred before that time. By 6000 B.P., west (mainland ) and to the rising sea levels had resulted in the formation north and east (the insular Pacific). Although of an island Southeast Asia from the ancient those authors traced one of those lines of land masses of and Sahulland descent east through the Bismarck Archi­ (Bellwood 1985; Green, in press). The con­ pelago and region to New sequent reduction in land area and hence pop­ Caledonia and Samoa (Schwarz and Schwarz ulation size of Rattus stock may have created 1967: 150), they derived the New Zealand a "bottleneck effect" leading to a speciation stock as follows (p. 151): "it is possible to trace event (Val 1988) from which R. exulans might ... that now stand as R. r. exulans Peale have been derived. from to the Caroline and Marshall Is­ Whatever the impetus for speciation, the lands; from there south to the Tonga Islands dispersal ofboth flora and fauna from South­ and New Zealand ..." (Figure 1). east Asia eastward into the Pacific has been Archaeological evidence ofthis rat from the impeded in regard to both space and time. postulated center of origin somewhere in is­ Reasons for this include the difficulty in cross­ land Southeast Asia comes from the Neolithic ing water, the distances between islands, and period, dating around 5000 B.P. the failure to find suitable habitats or chance associated with or immediately preceding following colonization. Biotic Neolithic pottery assemblages dated at 4100­ zones have been described on the basis of the 3700 B.P. in East Timor include the pig, observed inequities in species number and civet cat, macaque monkey, and R. exulans composition of the and plant biotas (Spriggs 1989). The emergence of R. exulans between islands in that region. For example, from ancestral Rattus stock must therefore "," lying between Borneo and Irian Origin, Dispersal, and Distribution of Rattus exulans-RoBERTs 125

10' /,t'J~~to~PHILLIPINES MARSHALL I

f/ j",. PALAU PONAPE ..... CAROLINE I >(. ::;,: MAJURO p.V~'··· ':.HO .: -- IIN BORNEO,' p.1.;\-•• ' - ~'~IIl:l:

KIRIBATI ·~~~LAWESI ~ c; N EAR - ~ l!...Nl:.. O' ~ : .?r.... '" 0 0 '" 0 C E A N I A c:> -_ '0J\rl BISMARCK' ~ Ntw Y (h'L- o ,'. <;> ~ RCHIPELAGO,",~IREL);N~ 'r", REMOTE NEW GUINEA • _ ...n. •3000 OCEANIA .~ ~ "- : FLORES "" () SOLOMON I ~ ~"- '\ ~Or;:!? ..c;:;=::>':/::::? c:,~ TUVALU . ~ U TIMOR SAN , . "4100 CHRISTOBAL'\> I '.sANTA CRUZ I 10' . / '. • TlKOPIA 1.290 ~ ., U\ ~FIJI o 0" 2700

NE~~Q. 20' CALEDONIA TONGA AUSTRALIA 2770

tlORFOLK I 8~900

KERMADEC I 30'

770 NEW ZEALAND '30' 150' 180' 170' W

FIGURE 2. Geographical locations ofthe "Thorne-Green" line and the "exulans-only" line within Near and . The radiocarbon dates (time Before Present) are the earliest estimates for Rat/us exulans.

Jaya (or more definitively, between the rodent genera occur there, including 16 species Wallace-Huxley line and Weber's line), marks ofRattus. This reduces to only three genera in a major faunal transition between the Oriental the , where three Rattus and Australian faunas. In fact, the Wallace­ species have been identified from prehistoric Huxley line (Figure 2) separates one of the sites in Balof, New Ireland (Allen et al. 1989) richest vertebrate faunas in the world from (i.e., R. mordax, R. praetor, and R. exulans, one ofthe poorest; beyond it to the east there the last-mentioned at around 3000 B.P. [Figure is a continuing trend toward faunal impover­ 2]). In fact, all nonvolant mammals other than ishment. This is true for both invertebrates Rattus sp. and Mus musculus reach the east­ (Laird 1956) and for vertebrates (Darlington ernmost limit of their distribution in the 1957). For instance, from New Guinea, 520 Solomon Islands (Thorne 1963). Pawley and species of birds have been recorded; the Green (1973) have delineated the eastern Solomons, 127; Fiji, 54; Samoa, 33; the boundary of this faunal distribution with a Society Islands, 17; the Marquesas, 11; and line passing between the southeastern end of Henderson, 4 (Thorne 1963). Of mammals, the Solomon Islands (San Cristobal) and the fOUf orders including the Rodentia are in­ Santa Cruz group 350 km to the east, thus digenous to New Guinea. Approximately 24 separating Near Oceania (New Guinea, the 126 PACIFIC SCIENCE, Volume 45, April 1991

Bismarck Archipelago, and the Solomon 3) confirm this association. Along with their Islands) from Remote Oceania (eastern pottery and animals, those people entered the Melanesia, , and Micronesia). They Fiji-Tonga-Samoa triangle sometime after noted that this line (referred to as the "Thorne­ 3500 B.P. but probably by 3000 B.P. (G. Irwin, Green" line in Figure 2) marks "a major cutoff pers. comm.). In that region, a cultural lineage point in the natural distribution ofanimal and was developed that subsequently dispersed plant species." Beyond this the only nonvolant farther east and whose descendants are mammals to proceed (by way of human as­ known today as Polynesians. On the far-flung sistance) were one or two rat species and the islands of Polynesia, the only indigenous domesticated dog, pig, and chicken. For ex­ mammals present (if any at all) were one or ample, the 2900-year-old mammalian fauna two species of bats. The rat, dog, pig, and ofTikopia on the Remote Oceania side ofthis chicken accompanied humans on those voy­ line includes the dog, pig, Rattus exulans, R. ages, but not all ofthose species survived each praetor, and two species offruit bats (Pteropus colonization event. For example, only the dog spp.), with only the bats likely to have been and rat apparently made it to New Zealand present before human colonization (Flannery (Pawley and Green 1973), but R. exulans is the et al. 1988). Approximating the "Thorne­ only vertebrate found in archaeological de­ Green" line but slightly farther to the east, a posits (dated at 800-900 B.P.) on Norfolk line can be drawn between the Santa Cruz Island (Meredith et al. 1985). Islands and Fiji indicating the boundary be­ Figure 3 summarizes these hypothesized yond which the only rat species known to have dispersal events through Near and Remote been introduced prehistorically is R. exulans. Oceania as outlined above. All dates are This line is labeled the "exulans-only" line in radiocarbon estimates of time of first human Figure 2. settlement (although the date for Easter Is­ Modern anthropological theory now ac­ land may be later than the time offirst settle­ cepts that the ancestors of the Polynesians ment). Some explanatory comments need to entered the Pacific by way of Melanesia be made. As the dates indicate, New Guinea, (Bellwood 1979). Their dispersal eastward the Bismarck Archipelago, and the northern from the Bismarck Archipelago to Tonga and Solomon Islands were all settled in the Pleisto­ Samoa between ca. 3600 and 3000 B.P. coin­ cene, but the rest of the insular Pacific was cides with the introduction of their com­ settled much later. The line linking Vanuatu mensals (the dog, pig, chicken, and rat) as has to the Marshall and Caroline islands is a been detailed above for New Ireland and language-based inference. The people ofwest­ Tikopia. The cultural complex ofthese partic­ ern Micronesian island groups (the Marianas, ular people has been called "Lapita" (Green Palau, Yap) all speak languages of Western 1979) in reference to the distinctive pottery Austronesian (island Southeast Asian) type, recovered from sites in the Bismarck Archi­ and thus they were presumably settled from pelago and the Solomon Islands east to Santa island Southest Asia. Eastern Micronesians Cruz, Vanuatu, New Caledonia and the (Caroline and Marshall islands, , Loyalty Islands, Fiji, and on to western Poly­ Tuvalu) speak Oceanic (i.e., Eastern Austro­ nesia (Tonga and Samoa). The archaeological nesian) languages. Those peoples thus seem to evidence suggests that the eastward dispersion be most closely related to those of northern of R. exulans across the biogeographical Vanuatu and the southeastern Solomon Is­ boundaries denoted by the "Thorne-Green" lands (Blust 1984-1985). The settlement dates and the "exulans-only" lines was made pos­ of the Cook and Society islands and the sible by the increasingly accomplished long­ Tuamotu Archipelago are unknown, but it is distance voyaging ofthese Lapita people. The very likely to be earlier than the settlement dates for R. exulans bones from Fiji (Best date of the Marquesas (ca. 2000 B.P.). Al­ 1984) and from Tonga (poulsen 1987) given though it is uncertain which particular island in Figure 2 allied with the dates for earliest group was first settled in central-eastern Poly­ human settlement in Remote Oceania (Figure nesia, the information in Figure 3 follows the Origin, Dispersal, and Distribution of Rattus exulans-RoBERTs 127

(l 1600 . HAWAUAN I 3000 MARIANA I ~HILLIPINES 20'

f' ~•.Jl . 2000 ?Q ". MARSHALL I

BISMARCK .~ :R;~.I~~~AGO 0'

If;) SOLOMON I , .. ~ " \a.. T1KOPJA 3000 2000 ~ 3000 SAMOA / MAROUESAS 0': --"""""3300 ~. /" VANUATU,. FIJI ~ .. SOCIETY I

3000~ '. <:J '~GA COOK ~ • ~

AUSTRALIA ,",7.~••. ' """:'/ ""'" . ~.~~;~.

NORFOLK 1./ ~ 1000 NEW ZEALAND r...... V o 2000k CH=~~AM '0' J1 :' 1

FIGURE 3. Reconstruction ofthe hypothesized insular Pacific dispersal routes ofRattus exulans, based on radiocar­ bon estimates (time Before Present) of time offirst human settlement. assumption that the island groups were settled by the (92 0 E long.) off the in order of accessibility from West Polynesia, west coast ofSoutheast Asia, while in the east according to recent theories of East Poly­ it has reached (109 0 W long.). nesian colonization (Irwin 1989). New Zea­ To the north it is found in Burma (25 0 N lat.) land, the last major world landmass to be and in (28 0 N lat.), while the southern­ settled by humans, was reached by ancestral most limit ofits distribution is New Zealand, Maori ca. 1000 B.P. (Irwin et al. 1990). The where it ranges as far south as Stewart Island earliest rodent evidence (presumably of R. (47 0 S lat.). Although it has been recorded exulans) in New Zealand comes from rat­ from two offshore islands of northern Aus­ gnawed berries of hinau (Elaeocarpus den­ tralia (Taylor and Horner 1973), that conti­ tatus) dated at 770 ± 50 B.P. recently dis­ nent remains the only major landmass in the covered at an archaeological site in Auckland Southeast Asia-Pacific region that has not (S. Best, pers. comm., 1990). been colonized by this species, possibly be­ cause its preferred grassland habitat was al­ Distribution ready occupied (Gressitt and Ziegler 1973). The complete geographical distribution Distribution within New Zealand of R. exulans is illustrated in Figure 4. The oceanic records shown here are detailed in Rattus exulans is known in New Zealand by Atkinson (1985). As can be seen, this rodent its Maori name, "kiore." Ethnographic ac­ has a fairly circumscribed distribution, sel­ counts of the arrival of this rat in New dom being found beyond the confines of the Zealand include those of the Maori people, tropics. Its westernmost boundary is marked whose oral histories have been recorded by 128 PACIFIC SCIENCE, Volume 45, April 1991

-rJ- - • . to -----~ -----~ ~ ---_.- --,----!~:~;e;f

.~sI i" .'....,,;.•, PAC I F I C 0 C E A N . .; ...... \. . • •• ~...... ,•• 1 •• •• f"'-...... • ~_,. .. .~- • - e.!!. • ~.... , .:• • 1. ... ". • _~ -- _a ! -' INDIAN OCEAN --• ., • --.... - ~ ).

FIGURE 4. Recorded world distribution of Rattus exulans. Reproduced by kind permission from Wodzicki and Taylor (1984). many authors. One of these, Best (1942), 1975). Earlier, in 1840, this species had be­ makes reference to a tradition that names come sufficiently scarce over much of the Aotea as the canoe into which R. exu/ans, the - North Island for Hutton (1879) to resort to sweet potato (Ipomoea batatas), swamp hen material collected from middens to describe (Porphyrio me/anotus), and seeds of the it as Mus maorium. The current distribu­ karaka tree (Corynocarpus /aevigata) were tion within New Zealand has been well doc­ placed before its departure from Hawaiki (the umented by Watson (1956), Taylor (1978), ancestral homeland). Other tribal traditions Atkinson (1986), and Atkinson and Moller give the name ofthis canoe as the Horouta and (1989). In brief, the only remaining mainland also refer to methods used by the Maori for populations ofR. exu/ans are in parts ofSouth trapping and cooking this rat, giving credence Westland and Fiordland in the South Island. to the possibility that its use as a supple­ However, many offshore island populations mentary food source led to its deliberate still persist, ranging from the Kermadec introduction and distribution. A. C. Ziegler Islands north of New Zealand (considered (pers. corom.) has recently suggested that part of territorial New Zealand) to Stewart these rats might also have served as a food and nearby islands in the south. On many of source for the domesticated dog, based on the the more northern offshore islands, R. exu/ans discovery of bone fragments of R. exu/ans is the only rodent species present, and, on a in the stomach region of a dog skeleton re­ number of these, it is also the only covered from a Hawaiian burial site. present. On a few of the larger islands it co­ After the introduction by Europeans of exists with one or more of the other rodent other rodent species (Rattus rattus, R. species, but nowhere in New Zealand do all norvegicus, and Mus museu/us) to New Zea­ four rodent species co-exist. After an ap­ land, there was a marked decline in the praisal of the historical decline of kiore on abundance of R. exu/ans on the main islands, New Zealand's mainland and an examination so that by the end of the nineteenth century it ofthe various combinations ofrodent species was considered virtually extinct (Taylor on the offshore islands, Taylor (1975, 1984) Origin, Dispersal, and Distribution of Rattus exulans-RoBERTs 129 suggested that strong competitive interactions LITERATURE CITED between species may be followed by displace­ ment and even exclusion. On New Zealand's ALLEN, J., C. GOSDEN, and J. P. WHITE. 1989. two main islands mice in particular seem to Human Pleistocene adaptions in the trop­ have been a major factor associated with the ical island Pacific: Recent evidence from disappearance of R. exulans. Such interac­ New Ireland, a Greater Australian outlier. tions and their effects are postulated to be Antiquity 63: 548-561. more severe in New Zealand than in tropical ATKINSON, I. A. E. 1985. The spread of com­ latitudes (where all four species can and do mensal species of Rattus to oceanic islands co-exist) because these islands are at the ex­ and their effects on island avifaunas. Pages treme southern limits of this originally trop­ 35-81 in P. J. Moors, ed. Conservation of ical rat's range (Taylor 1975). island birds. I.C.B.P. Tech. Pub!. no. 3. Cambridge, England. ---. 1986. on New Zealand's CONCLUSIONS northern offshore islands: Distribution, effects and precautions against further The ongm of R. exulans from ancestral spread. Pages 13-40 in A. E. Wright and stock somewhere in island Southeast Asia, as R. E. Beever, eds. The offshore islands of postulated by Tate (1935) and Schwarz and northern New Zealand. Department of Schwarz (1967), remains unchallenged. How­ Lands and Survey Information Series no. ever, current anthropological and archaeo­ 16. Wellington, New Zealand. logical information allow a reconstruction of ATKINSON, I. A. E., and H. MOLLER. 1989. their views concerning the probable historical Kiore, Rattus exulans (Peale). Pages 175­ dispersal route of R. exulans throughout the 192 in C. King, ed. Handbook ofNew Zea­ insular Pacific and, in particular, to New Zea­ land mammals. Oxford University Press, land. This modern hypothesis, outlined in Auckland, New Zealand. Figure 3, differs from that of earlier authors BELLWOOD, P. 1979. The oceanic context. (Figure 1) by deriving the New Zealand stock Pages 6-26 in J. D. Jennings, ed. The pre­ not from a migration line passing through history of Polynesia. Harvard University Micronesia, but from one that passes through Press, Cambridge, Massachusetts. Near to eastern Remote Oceania and thence ---. 1985. Pre-history of the Indo­ south to New Zealand. In addition to updat­ Malaysian Archipelago. Academic Press, ing theories concerning the dispersal routes of Sydney, Australia. R. exulans in the Pacific, Figure 2 presents BEST, E. 1942. The forest lore of the Maori. radiocarbon dates for the estimated arrival Dom. Mus. Bull. 14. times of R. exulans in Near Oceania and BEST, S. B. 1984. : Prehistory of a western Polynesia. The figure is intended to Fijian Island. Ph.D. thesis, University of stimulate interest in the accuracy (or other­ Auckland, Auckland, New Zealand. wise) of the "exulans-only" line and to elicit BLUST, R. 1984-1985. The Austronesian more information relating to the prehistory of homeland: A linguistic perspective. Asian this species elsewhere in the insular Pacific. Perspect. 26 :45-67. BUCK, P. 1938. Vikings of the sunrise. J. B. Lippincott. Reprint, 1958, Whitcombe and Tombs, Christchurch, New Zealand. ACKNOWLEDGMENTS DARLINGTON, P. J. 1957. Zoogeography; the I am indebted to staffofthe Department of geographical distribution of mammals. Anthropology at Auckland University; in Wiley, New York. particular, I would like to thank Geof. Irwin, FLANNERY, T. F., P. V. KIRCH, J. SPECHT, and Roger Green, Richard Walter, Dorothy M. SPRIGGS. 1988. Holocene mammal Brown, and Liz Hudson for their helpful dis­ faunas from archaeological sites in island cussions, assistance, and encouragement. Melanesia. Arch. Oceania 23 :89-94. 130 PACIFIC SCIENCE, Volume 45, April 1991

GREEN, R. C. 1979. Lapita. Pages 27-60 in SPRIGGS, M. 1989. The dating of the Island J. D. Jennings, ed. The prehistory of Poly­ Southeast Asian Neolithic; an attempt at nesia. Harvard University Press, Cam­ chronometric hygiene and linguistic cor­ bridge, Massachusetts. relation. Antiquity 63: 587-613. ---. (in press). Near and Remote TATE, G. H. H. 1935. Rodents of the genera Oceania-disestablishing "Melanesia" in Rattus and Mus from the Pacific Islands, culture history. In A. K. Pawley, ed. Man collected by the Whitney South Sea Expedi­ and a half: Essays in Pacific anthropology tion, with a discussion of the origins and and ethnobiology in honour of Ralph races of the Pacific Island rat. Bull. Am. Bulmer. Auckland University, Auckland, Mus. Nat. Hist. 68: 145-178. New Zealand. TAYLOR, J. M., and B. E. HORNER. 1973. Re­ GRESSITT, J. L., and A. C. ZIEGLER. 1973. The sults of the Archbold Expeditions. No. 98. effect of the loss of forests in New Guinea. Systematics of native Australian Rattus Pages 117-122 in A. B. Costin and R. H. (Rodentia, ). Bull. Am. Mus. Nat. Groves, eds. Nature conservation in the Hist. 150: 1-130. Pacific. Proc. Symp. 12th Pac. Sci. Congr., TAYLOR, R. H. 1975. What limits kiore Canberra, 1971. Australian National Uni­ (Rattus exulans) distribution in New Zea­ versity Press, Canberra, Australia. land? N. Z. J. Zool. 2(4): 473-477. HUTTON, F. 1879. Note accompanying speci­ ---. 1978. Distribution and interactions mens of the (Mus rattus). Trans. of rodent species in New Zealand. Pages N.Z. Inst. 11: 343-344. 135-143 in P. R. Dingwall, 1. A. E. IRWIN, G. 1989. Against, across and down the Atkinson, and C. Hay, eds. The ecology wind; a case for the systematic exploration and control of rodents in New Zealand of the remote Pacific Islands. J. Polynesian nature reserves. Department of Lands and Soc. 98(2): 167-206. Survey Information Series no. 4. Welling­ IRWIN, G., S. BICKLER, and P. QUIRKE. 1990. ton, New Zealand. Voyaging by canoe and computer: Experi­ ---. 1984. Distribution and interaction of ments in the settlement of the Pacific introduced rodents and carnivores in New Ocean. Antiquity 64: 34-50. Zealand. Acta Zool. Fenn. 172: 103-105. LAIRD, M. 1956. Studies on mosquitoes and THORNE, R. F. 1963. Biotic distribution pat­ freshwater ecology in the South Pacific. R. terns in the Tropical Pacific. Pages 311-354 Soc. N.Z. Bull. 6: 192-194. in J. L. Gressitt, ed. Pacific Basin biogeog­ MEREDITH, C. W., J. R. SPECHT, and P. V. raphy. Bishop Museum Press, Honolulu, RICH. 1985. A minimum date for Poly­ Hawaii. nesian visitation to Norfolk Island, South­ VAL, F. C. DO. 1988. Speciation in the Neo­ west Pacific, from faunal evidence. Search tropics and the founder principle. Pac. Sci. 16(9-12): 304-306. 42: 105-114. PAWLEY, A., and R. GREEN. 1973. Dating the WATSON, J. S. 1956. The present distribution dispersal ofthe Oceanic languages. Oceanic and abundance of Rattus exulans (Peale) in Linguistics 12: 1-67. New Zealand. N. Z. J. Sci. Tech. 37: 560­ POULSEN, J. 1987. Early Tongan prehistory: 570. The Lapita period on Tongatapu and its WODZICKI, K., and R. H. TAYLOR. 1984. Dis­ relationships. Terra Australis 12, vol. I: 1­ tribution and status of the Polynesian rat 307. Rattus exulans. Acta Zool. Fenn. 172: 99­ SCHWARZ, E., and H. K. SCHWARZ. 1967. A 101. monograph of the Rattus rattus group. Ann. Esc. Nac. Cienc. BioI., Mexico City 14: 79-178.