THE PRE-EUROPEAN AND PRESENT DISTRIBUTIONS OF GODMAN! (MARSUPIALIA: ), A RESTRICTED RAINFOREST ENDEMIC

WILLIAM F. LAURANCE

Laurance, W. F., 1993. The pre-European and present distributions of Antechinus godmani (Marsupialia: Dasyuridae), a restricted rainforest endemic. Australian Mammalogy 16(1): 23-27.

New capture localities for the Atherton Antechinus (Antechinus godmani), a restricted endemic in north Queensland rainforest, help clarify the northern limits of its geographic range. I summarise these records, then use recent knowledge of the habitat and altitudinal requirements of A. godmani to estimate both its pre-European and present distributions, as well as the impact of deforestation on this species. Before 2 European settlement, the total range of A. godmani may have encompassed an area of 2900 km . Since 2 then, its range has been reduced by more than 820 krn , resulting in extensive fragmentation of populations 2 in the northern half of its range. The current distribution of A. godmani may exceed 2000 km , a figure that easily triples earlier estimates for this species.

Key words: Antechinus godmani, deforestation, geographic range, north Queensland, rainforest endemic.

William F. Laurance, MuseumofVertebrate Zoology, University of California, Berkeley, CA 94720, USA. Current address: Wet Tropics Management Agency, PO Box 2050, Cairns, Queensland, 4870. Manuscript received 12 December 1991.

THE Atherton Antechinus (Antechinus godmani) is forests had reduced and subdivided the species into a small (< 130 g) marsupial predator known only from smaller subpopulations. McDonald (1991) described upland rainforest in the Atherton region in north ten capture sites for A . godmani in the Cardwell and Queensland. Considered rare and poorly known, its Kirrama Ranges located up to 55 km south of the geo:rraphic range was initially estimated at only 600 previously known range of the species. km giving it one of the smallest known distributions Herein, I summarise documented capture localities of any Australian species (VanDyck 1983). for A. godmani and describe nine new or unpublished VanDyck (1982) suggested that A . godmani should capture sites that help clarify the northern limits of its be considered a "relict endemic" of tropical geographic range. I use recent knowledge of the Queensland, because of its restricted range, habitat and altitudinal requirements of A. godmani to conservative morphology, and lack of vicariant sister estimate both its historical (pre-European) and current groups (Kikkawa, Monteith and Ingram 1981). distributions, as well as the impact of deforestation on Winter, Bell, Pahl and Atherton (1984) used the this species. known range of A . godmani to define the centre of a 'refugial area' in north Queensland, and demonstrated MATERIALS AND METHODS that this region supports an exceptionally diverse assemblage of non-flying . NEW DISTRIBUTIONAL RECORDS Table I lists all confirmed capture localities for Before 1990, A. godmani was reported at only six A. godmani. Details of the new records and their localities, all located within 30 km of the town of localities (circa June 1992) follow. Ravenshoe on the southern Atherton Tableland (Van Dyck 1982). However, since then there has been a On 26 March 1991, I captured an adult male spate of new records that has increased the known A. godmani (testes scrotal, weight = 84 g) in range of A . godmani. Laurance (1990a) described continuous, complex mesophyll vine-forest (Tracey three new localities for A. godmani on the south-east and Webb 1975) on the north-east margin of the margin of the Atherton Tableland, and argued that Atherton Tableland (10.0 km E, 5.5 km N of clearfelling of suitable upper elevation (above 600 m) Yungaburra; l7°!3'05"S, 145°41 '15"E) at 780 m 24 AUSTRALIAN MAMMALOGY elevation. The trap was positioned 10m from the verge L.A. Moore and F. H. J. Crome (pers. comm.) report of the Gillies Highway at a site overlaying granitic captures of 17 individuals of A. godmani at four sites soils. near Butcher's Creek on the eastern margin of the Atherton Tableland (1. 7, 3.3, 4.7, and 7.8 km E of A specimen collected and preserved by J. W. Winter Larnins Hill, respectively; Table 1), all in complex (pers. comm.) on 23 October 1974, was positively mesophyll vi ne-forest between 750--780 m elevation. identified recently as A. godmani by S. Van Dyck. The Significantly, they further report capturing two , a small fema le, was captured at 1530 m in an isolated, 190 ha fragment of complex elevation near the summit of Mt Bellenden Ker notophyll vine-forest located on the central Atherton (0.6 km SW of the Centre Peak; l7°!5'45"S, Tableland (2.0 km N, 0.5 km E of Tarzali; 17° 24' 145°51' 15"E). This locality is the highest elevational 26" S, 145° 36' 23 " E) at 760 m elevation. record for the species, the first capture in rnicrophyll vine-forest, and the only record from the Mt Bellenden Finally, interrogation of the mammal database from Ker vicinity. the Queensland Museum (S. Van Dyck, pers. comm.) revealed three unpublished (1989-1990) capture localities for A. godmani, one of which (Record No. JM 8031) is presently the northernmost record for the Latitude Longitude Elevation Year Source (m) species (Danbulla State Forest; l7°06'S, 145°38'E). All confirmed localities from current Queensland 17"42' 145.31' 1921 I Museum records (June 1992), excepting four early 1TIS.7S' 145.51.25' 1520-1540 1974 2 (1921 - 1969) localities which were described 17"44' 145.32' 780-850 1976 I imprecisely, are summarised in Table 1. These records 1T36' 145.39' 710 1976 I have been vetted carefully and should supersede 145.39' 1977 I 17"41' localities given in VanDyck (1982) (S . Van Dyck, 17"40' 145.39' 1977 I 17"24' 145.41' 700 1981 3 pers. comm.). 17"44' 145.33' 1982 I 145.37.67' 1987 4 17"32.33' 720 CRITERIA FOR EsTIMATING GEOGRAPHIC IT36.33' 145.37.75' 800 1987 4 17"28.17' 145.31.25' 600 1987 4 RANGES 18.01.83' 145.36.83' 980-1000 1989 s I estimated the historical and current ranges of 18.02.92' 145.36.33' 760-780 1989 s 18. 13.08' 145.47.42' 800-820 1989 s A. godmani by using maps from the Australian 18. 14.16' 145.47.75' 880-900 1989 s Topographic Survey (I : 100 000 map numbers 7962, 18. 15.28' 145.47.97' 840-860 1989 s 7963, 8063, circa 1970; and 8061, 8062, circa 1980; 18.15.30' 145.47.92' 880-890 1989 s 1:50 000 map numbers 806 1-1, 8061-2, 8061-4, 18. 15.22' 145.48.08' 880-900 1989 s 8064--3; circa 1980), augmented with infrared Landsat 18. 15.33' 145.48.33' 880-900 1989 s imagery (circa 1981 ), and local knowledge. 18. 16.83' 145.48.58' 800-820 1989 s Throughout the analysis I worked on the scale of 18. 17.92' 145.48.83' 800-820 1989 s I km2 blocks; for example, if a particular block !TIS' 145.40' 680 1990 I occurred on a rainforest-sclerophyll forest margin, I 17"06' 145.38' 760 1990 I 17"24.43' 145.36.38' 760 1990 6 included it in the analysis if 50 % of the block was 17"22.62' 145.43.33' 780 1990 6 covered by rai nforest. 17.22.58' 145.43.75' 760 1990 6 17"22.70' 145.44.53' 750 1990 6 Antechinus godmani has been captured in all major 17"21.80' 145.45.83' 760 1990 6 rainforest and soil types in the Atherton Uplands (Bell, 17"13.08' 145.41.25' 780 1991 7 Winter, Pahl and Atherton 1987), including selectively logged areas, but never below 600 m Table I. Confirmed capture sites for Antechinus godmani elevation (Laurance 1990a; McDonald 1991). Rather in north Queensland rainforest, as at June 1992. Latitude than being restricted to certain rainforest types, and longitude data are judged to be accurate to within one available evidence suggests its range is limited by minute or less ( < 1.8 krn) except for the type locality major low-altitude features such as river gorges (recorded in 1921 ), which is considered accurate to within (McDonald 1991 ). five minutes or less (< 9 km). Four early (1921 -1969) capture localities that were imprecisely described are not The historical range of A. godmani was estimated included. "Year" indicates the date of the first recorded by pooling: (I) all sizeable(> 200 ha) rainforest tracts capture at each locality. Sources are: 1. Queensland Mu- within its probable elevational range (600--1685 m) seum Records, circa 1992 (S . VanDyck, pers. comm.); 2. J. W. Winter, pers. comm.; 3. Preen 1981; 4. Laurance that currently are contiguous, or nearly contiguous, 1990a; S. McDonald 1991; 6. L. A. Moore and F. H. J. with the Atherton Uplands. (Existing areas of Crome, pers. comm.; 7. W. F. Laurance, pers. obs. microphyll vine-forest, such as those ranging up to LAURANCE: DISTRIBUTION OF ANTECHINUS GODMAN/ 25

1685 m elevation at the summit of Mt Bartle Frere are lower-elevation(< 400 m) forests associated with the included, because A. godmani has been captured in Herbert River to the south (McDonald 1991) and the this forest type); (2) extensive areas of the Atherton Barron River to the north. Uplands that have been deforested or inundated by The mapping exercise has several merits in addition artificial reservoirs since European settlement; and to estimating the historical and current ranges of (3) rainforest tracts such as the Herberton Range and A. godmani. Hugh Nelson Range which, although isolated from the Atherton Uplands, occur within the appropriate The exercise highlights several poorly studied elevational range (600-1685 m) and obviously were areas, such as Mt Fisher and Mt Bartle Frere, the linked with the Atherton Uplands prior to Herberton and Hugh Nelson ranges, and the eastern deforestation. Walter Hill Range, that are predicted to support The estimate of historical range excluded: (I) natural lakes; (2) an area of I 04 km2 on the drier north-west corner of the Atherton Tableland that probably was dominated by wet sclerophyll forest prior to European settlement (see dotted line, Fig. I); r and (3) disjunct mountain peaks separated from the N Atherton Uplands by river gorges or by large expanses 2 I (5 km ) of low-elevation (< 300 m) forest, which probably would have acted as barriers to dispersal by A. godmani. To estimate the amount of range of A. godmani that has been rendered unusable since European settlement, I pooled: (I) deforested lands within the appropriate elevational range, again excluding 104 km2 on the drier north-west corner of the Tableland; (2) two artificial reservoirs (Koombooloomba and Tinaroo dams); (3) large (>I km) pine plantations; and (4) isolated rainforest fragments of <100 ha in area, because A. godmani apparently does not survive in these areas (Laurance 1989, 1990a, 1991). RESULTS AND DISCUSSION O Km PAST AND PRESENT RANGES The historical range of A ..podmani is estimated to have encompassed 2898 km (Fig. I). However, since 2 European settlement began in the late 1800s, 822 km , or 28.4% of the original range of A. godmani, has been destroyed or rendered unusable. This leaves a total of 2076 km2 of potential habitat remaining, a fifure that easily triples the earlier estimate of 600 km for this species (VanDyck 1983). CJ Antechinus godmani appears to be limited to the narrow band ( < 30 km wide) of upland rainforest Fig. / . The distribution of Antechinus godmani in the above 600 m elevation south of Cai rns in far north Atherton Uplands region of far north Queensland. Filled Queensland. The latitudinal range of the species, circles denote new or undocumented capture localities for the based on current capture records, spans a di stance of species; open circles are previously reported localities. 131 km from I7°06'S to I8°18'S. However, I suggest Darkly stippled areas are predicted tracts of suitable habitat for A. godmani (upland rainforest between 600-1685 m that the species could occur between 16°53' to 2 elevation, totaling 2076 km in area). Unstippled areas were !8°24' S, a latitudinal range of 166 km. This prediction deforested during the last century; irregular black areas are is based on the distribution of existing, apparently inundated by man-made reservoirs. Diagonally hatched areas suitable rainfo rest in the Atherton Uplands region to the east are rainforest below the 600 m elevationallimit of (Fig. l ). Ultimately, the latitudinal range of the species A. godmani, whereas lightly stippled areas to the west are may be limited by the gorges, floodplains and drier forests or woodlands. 26 AUSTRALIAN MAMMALOGY

A. godmani but for which there have been no captured an A. godmani in 1991 (17°13'05"S, documented records of this species. 145°41' IS"E). (3) The BIOCLIM map also did not include the northern half of the Atherton Tableland, It demonstrates also that patterns of forest clearing which now is mostly deforested, whereas my analysis on the Atherton and Evelyn Tablelands have had an suggests that all but the drier NW portion of the important impact on A. godmani. For example, it is tableland was formerly suitable habitat. Although this now evident that much of the northern half ofthe range difference is difficult to resolve because of extensive of A. godmani has been deforested, probably deforestation, my study suggests that larger(> I 00 ha) fragmenting the species into several discrete forest fragments on the northern Atherton Tableland, subpopulations (Laurance 1990a). such as Lake Eacham and Lake Barrine national parks The mapping exercise also highlights the and Wongabel State Forest, could support relict conservation significance of the narrow band of populations of A. godmani. (4) The BIOCLIM map suitable, upper elevation forest(> 600 m) fringing the predicted that upper-elevation (600-1685 m) forests eastern margin of the Atherton Tableland, some of associated with the large granite massifs of Mts which is freehold land potentially subject to Bellenden Ker and Bartle Frere provide only marginal deforestation. habitat for A. godmani, whereas my map suggests these areas are suitable habitat. (5) The BIOCLIM map The study may have significant biogeographic suggests that A. godmani will occur only on disjunct implications, because the restricted range of mountaintops in the Cardwell Range near the southern A. godmani has been used to identify a key centre of limit of its range, whereas my map suggests a larger mammalian endemism on the southern Atherton and more continuous distribution. Tableland (Winter et al 1984), which presumably resulted from the area's persistence as a Pleistocene MICROHABITAT SELECTION forest refuge (Laurance 1987; Winter 1988). Available evidence suggests that A. godmani is CLIMATE AS A POSSIBLE LIMITING FACTOR entirely rainforest-dependent. The species has been captured in larger rainforest fragments on the Atherton Recently, Nix and Switzer (1991) used a BIOCLIM Tableland ( 190 ha and 590 ha in area; Laurance 1990a; computer analysis to predict areas of suitable habitat L.A. Moore and F.H.J. Crome, pers. comm.) but there for endemic rainforest vertebrates in north have been no reported captures in pastures or other Queensland, including A. godmani. Their analysis non-forested habitats (Laurance 1989). produced a bioclimatic profile for each species, which was then mapped, producing a predicted distribution. Even within areas of suitable habitat, A. godmani The BIOCLIM map did not include effects of appears to be rare and patchily distributed. For deforestation or area-based estimates of the historical example, Laurance (1989, 1990a, in press) captured and current ranges of A. godmani. However, because A. godmani at only three of 20 sites in continuous the BIOCLIM map was produced under a different set forest on the southern Atherton Tableland, whereas of assumptions than mine (ie that climatic regimes or McDonald (1991) captured A. godmani at 10 of 31 their correlates, rather than geographic barriers, limit sites in the Kirrama and Cardwell Ranges. Although the range of A. godmani), it is useful to contrast it with the type of bait used and time of year influence capture my estimate of historical range. success (Laurance 1992), A. godmani appear to favour sheltered locations (McDonald 1991) with Although roughly concordant, the two predicted dense ground cover (Watt 1992; L.A. Moore, pers. ranges differed in several important ways: (I) The comm.). These microhabitats may J·rovide additional BIOCLIM map included two upland tracts, Mt substrates for foraging and a more consistent supply Carbine Tableland and Thornton Peak, located of insects than exposed areas subject to greater 30-80 km north of the Barron River, whereas I edaphic variation (for example, Frith and Frith 1985). identified the Barron River as the northern limit for Species such as A. godmani that are patchy, rare, and A. godmani. Although no specimens have been entirely rainforest-dependent typically will have recorded at either northern locality, these areas small, isolated populations in fragments, and hence obviously merit additional census work, particularly should be particularly vulnerable to habitat as several other upland rainforest mammals (for fragmentation (Terborgh and Winter 1980; Laurance example, Hemibelideus lemuroides, Pseudocheirops 1990b, 1991). archeri, Antechinus stuartii, Melomys hadrourus; F. H. J. Crome, pers. comm.) have distributions that The extent to which a species is arboreal is span the Barron River. (2) Unlike my analysis, the important because deforestation, as well as linear BIOCLIM map failed to include upland (> 600 m barriers such as roads, highways and powerline elevation) forests fringing the north-eastern margin of corridors, are more likely to act as physical barriers to the Atherton Tableland, including the site where I arboreal animals. Antechinus godmani is often LAURANCE: DISTRIBUTION OF ANTECHINUS GODMAN/ 27

captured in traps set on the ground (for example, LAURANCE, W. F., 1990a. Distributional records for two Laurance 1992), indicating that the species is at least "relict" dasyurid in north Queensland partially scansorial and hence should be physically rainforest. Australian Mammalogy 13:215-218. capable of crossing roads. Watt (1992) found that LAURANCE, W. F., 1990b. Comparative responses of five A. godmani fitted with radio-tracking devices readily arboreal marsupials to tropical forest fragmentation. crossed narrow logging tracks (< 5 m wide), but Journal of Mammalogy11 : 641-{)53. crossed larger roads and clearings (10-20 m wide) LAURANCE, W. F. , 1991. Ecological correlates of extinction much less frequently, especially if canopy cover was proneness in Australian tropical rain forest mammals. removed. Conservation Biology 5: 79-89. Clearly, further study is needed to determine whether LAURANCE, W. F., 1992. Abundance estimates of small larger disjunctions such as highways or powerline mammals in Australian tropical rainforest: a comparison clearings present a psychological barrier to movements of four trapping methods. Wildlife Research 19: of individuals (for example, Burnett 1991). 651-655. In summary, on one hand, the mapping analysis is LAURANCE, W. F., (in press). Rainforest fragmentation and the structure of small mammal communities in tropical cause for some optimism, as it suggests that the current Queensland. Biological Conservation. geographic range of A. godmani could exceed 2 2000 km , an area considerably larger than formerly McDONALD, K. R., 1991. New distributional records for proposed (Van Dyck 1983). On the other hand, the Antechinus godmani (Thomas), a restricted rainforest endemic. Memoirs of the Queensland Museum 30: analysis suggests that large-scale deforestation on the 487-491. Atherton and Evelyn Tablelands, as well as forest inundation by artificial reservoirs, have reduced the NIX, H. A. AND SwnzER, M. A., 1991. Rainforest animals: historical range of A. godmani by about 30 %, atlas of vertebrates endemic to Australia's wet tropics. extensively fragmenting populations in the northern 1, Australian National Parks and Wildlife Service: Canberra. half of its range. PREEN, A. R., 1981. The effects of selective logging on the ACKNOWLEDGEMENTS vertebrate fauna of a tropical rainforest in north-east Queensland. B.Sc. Hons. thesis, James Cook University: J. W. Winter, F. H. J. Crome, K. R. McDonald, and S. Townsville, Queensland. M. Van Dyck commented on an earlier draft of the TERBORGH, J. AND WINTER, B., 1980. Some causes of manuscript.lthankespecially L.A. Moore, S .. M. Van extinction. Pp. 119-133 in Conservation biology: an Dyck, F. Crome, A. Watt, and J. Winter for access to evolutionary-ecological perspective ed by M. E. Soule unpublished data, and K. Means for useful discussion. and B. A. Wilcox. Sinauer Associates: Sunderland, Assistance in the field was provided by students at the Massachusetts. S.F.S. Centre for Rainforest Studies, Yungaburra, TRACEY, J. G. AND WEBB, L. J., 1975. Vegetation ofthe humid Queensland. tropical region of north Queensland (1 :100 000 maps and key). CSIRO Long Pocket Labs: lndooroopilly, REFERENCES Queensland. BELL, F. C., WINTER, J. W., PAHL, L. 1., AND ATHERTON, R. VAN DYCK, S., 1982. The status and relationships of the G. 1987. Distribution, area and tenure of rainforest in Atherton antechinus, Antechinus godmani (Marsupialia: northeastern Australia. Proceedings ofthe Royal Society Dasyuridae). Australian Mammalogy 5: 195-210. ofQueensland98: 27-39. VAN DYCK, S., 1983. Atherton Antechinus. Pp. 43 in The BuRNETT, S., 1991. Effects of a rainforest road on movements Australian Museum complete book of Australian of small mammals: mechanisms and implications. 1nammals ed by R. Strahan. Angus and Robertson: Wildlife Research 19: 95-104. Sydney. FRITH, C. B. AND FRITH, D. W., 1985. Seasonality of insect WATT, A. M., 1992. The ecology of three species of abundance in an Australian upland tropical rainforest. Antechinus (Marsupialia: Dasyuridae) in upland Australian Journal of Ecology 10: 237-248. rainforest in north-east Queensland. Ph.D. thesis, James KIKKAWA, J., MONTEITH, G. B., AND INGRAM, J. M., 1981. Cook University: Townsville, Queensland. Cape York Peninsula: major region of faunal WINTER, J. W., 1988. Ecological specialisation of mammals interchange. Pp. 1697-1742 in Ecological biogeography in Australian tropical and sub-tropical rainforest: in Australia, ed by A. Keast. Dr. W. Junk: The Hague. refugial or ecological determinism? Proceedings of the LAURANCE, W. F., 1987. The Rainforest Fragmentation Ecological Society of Australia 15: 127-138. Project. Liane 25: 9-12. WINTER, J. W., BELL, F. C., PAHL, L. 1. , and ATHERTON, R. LAURANCE, W. F. , 1989. 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