Paper.\ and Proceedings of !lie !
NOTES ON THE BIOGEOGRAPHY OF THE FREE-LIVING TERRESTRIAL INVERTEBRATE FAUNA OF MACQUARIE ISLAND WITH AN ANNOTATED CHECKLIST
I\·nelupc Cireer1slade
(with six tables and an appendix) GREENSLADE, P., 1990 (3 l :x ): Notes on the biugeography of the frec-l1ving tcrre,tnal rnvcrtcbrnte la1111a of Macquarie Island with an annotated checklist. Pap Proc. R. Soc Tasm. 124(1): 35-50. https://doi.org/10.26749/rstpp.124.1.35 ISSN 0080-4703. CSIRO Division of Entomology, GPO Box 1700, Canberra. ACT Australia 2601. The invcrlehr,tte fauna of Mac4uarie bland is examillcd and it� siD.:, cotnposirion c-nlktnkily a11cJ biogeugt aph!cal affinit1c� are l·on1pared with tho.-..;c- of other �ubantarctic island�. Its origin.-;, dispersal mecha11hrn� and adapt ;itinw.: !!TC di,cussed and !he effect on it of vertebrate predators considered. A chccklisl ut the invcrtcbrnte.s known from Macquarie Island incorporating recent revisionary taxonomy is included as an appendix. Key Words: Subantarctic, Insecta, Collembola, Mollusca, Oligochaeta, Arachnida, endcmism.
INTRODUCTION Neither are the Acari listed here in detail. Womersley ( 1937a) published some of the earliest Prior to fieldwork carried out in the summer of records of mites and, although good collections are 1986-87 by the author. the only large collection of available from Macquarie Island, the faunas of terrestrial invertebrates from Macquarie Island was adjacent territories have nut been intensively that of Watson (1967) who recorded 144 species, of studil.'.d, su that a1,y list here would give a which he believed 49 (34%) were endemic. A later rnisleadrngly elevated for the percentage work by Cressit (1970) added little further endemism in this group (i.e. Wallwork 1973). Over taxonomic information, although, for the first time, 80 species of mites have been identified (D. the invertebrate fauna of Macquarie Island was Horning. pers. cornm.J, but the fauna certainly compared in detail with those'of other subantarctic contains 111ore than this and no revisionary work has islands. Since then, revisionary taxonomy of a been carried out for about 20 years. For this reason number of taxa has been published on the basis of they have not been included in the discussion. more exten,ive material from throughout the ·· Adventttious" species, which include those which subantarctic region (Selkirk et al. 1986). Using new disperse from time to time to the island but are information, a revised list has been compiled of unable to establish permanent populations, i.e. species known from Macquarie Island (see naturalise, v1ere li�ted hy Watson ( l 967) and are not appendix); this includes a number of new records, A repeated here. The ectoparasitic Phthiraptera will he trend observed, during its compilation, was a steady discussed in a later publication (D. Horning & R. reduction in percentage endcmism from the 1967 Palma, pers. comm.). (Watson) and l 970 (Gressitt) figures. The level of Tax a are arranged as in lnsee/s of Australia endemism f'or Macquarie Island is now much lower (CSIRO 1970) and Grccnsladc ( 1985 ). Full syn than for other suhantarctic islands and possible onymies are not given for the species which occur reasons fur this are discussed here. widely outside Macquarie Island but most names This new checklist excludes a number uf the lower mentioned in publication., on Macquarie Island arl: invertebrate taxa which arc inadequately collected included, except for th use of C'olknlilola which have and docs not covt.T fre,shwater species. of which recently been given by Greens lade & Wisc (1986). there are vc,ry few. l·or example, although species of Distribution records of each spec1t:s arc included Protozoa (Walson 1967), Rotifern (Watson 196'/). and arc classified according to whether they Turbcllaria (Ball & Hay 1977), Nematoda (Bunt have hccn introduced relatively recently, probably 1954 ), Copcpoda (Hamond 1987) and Cladocera hy humans, or arc considered native to the island. (Smirnov & Timms l 983) have been recorded from The methud used to determine to which category the island. l!1ost of these group,s arc still poorly each species belongs has heen described by known, All species so far recorded in them, apart Greemlade ( I 'J87a). Material collected in l 986--87 from the Turbellaria, have widespread distributions. is deposited either in the South Australian Museum
35 36 Greenslade
(SAlVIA) (Collembola) or in the Australian Nalional 1(86). Twenty-six insect have been recorded Insect Collection (ANIC) (all other invertebrates). from Marion Island compared 'vvith 31 from Fieldwork was conducted in December 1986 and Macquarie Island (Table 4). January J 987. Samples came from as many different Returning to the Collembola, the size of the habitats and sites on the island as possible and over Macquarie Island fauna j, of the same order ,,1' 100 collections were made. Methods used included magnitude as those of rhe and Crozet pitfall traps, funnel extraction of soil and leaf litter, Islands (Deharveng 198]) but if nattve species sweeping and beating vegetation and are included, it is smaller CrabJe 5). Marion Island under stones. A preliminary version of this checklist has a smaller fauna but it is undercollected was published in Selkirk et al. (1990). while Heard Island, which has little ice-free area and then for only a short period in the summer, has an even smaller fauna (Greenslade 1986). Until CI1ARACTERISTICS OF THE Collembola have been collected from the other MACQUARIE ISLAND FAUNA suhantarctic islands as as from Macquarie Island, further comment cannot be made Size about relative size of faunas.
So far as number of invertebrate species is concerned it is clear that the fauna is extremely Composition small for a humid vegetated island of area 12800 ha (120 km2, 34 x 2.5-5 km). Only 76 resident species The representation of higher taxa on Macquarie of terrestrial arthropods, molluscs and oligochaetes Island is very uneven. There are as many coHem (excluding mites) (Table 1) are recorded despite bolan as insect species. Among the insects, the considerable coilecting effort. Although some Diptera are the best represented group but only two Diptera species are yet to be identified, it is unlikely families of Coleoptera are present and among the that this list will be increased by much. other large orders, there are few Hemiptera, Taking a single group, the Collembola, of which Lepidoptera and Hymenoptera. 31 species are listed, five were found in the The composition of the insect fauna is similar to greenhouse among the main station buildings. In that of Marion Island in that five of the same orders contrast, one can generally find 30 species of of native insects occur. The main differences lie in Collembola on a site of 10 m 2 in southern Australia. that there are no Curculionidae, fewer Lepidoptera Even on Philip Island, a much smaller island (3 km2) but more Staphylinidae and Diptera species on off the coast of Norfolk Island in the Tasman Sea, lVIacquarie Island Cfable 4) compared with Marion and at that time practically denuded of vegetation, Island. 25 species have been collected (P. Greenslade), The composition of collembolan faunas, so far as while The Snares, an undisturbed island group north genera are concerned, is similar on all the sub of Auckland Island, simiiar in size to Philip Island, antarctic islands; genera such as Cryplopygus and carries nearly 50 species of Collembola (P. Tullhergia are the most diverse and are character Greenslade). istic of these faunas. The Kerguelen Islands have the A comparison of the faunal size with that of other largest and most varied collembolan fauna in terms subantarctic islands is difficult because there are of both genera and (Deharveng 1981). many variables in terms of latitude, area, climate, altitude, permanent snow cover and origin. (The term subantarctic islands is used here in Endemism its strict sense and includes only the Kerguelen, South Georgia, Heard, Prince Edw(lrd group A level of endemism (34°l), was recorded for (including Marion Island), Crozet and Macquarie Macquarie Island invertebrates by Watson (1967); Islands.) The most similar island to Macquarie when mites were excluded, this increased to about Island appears to be Marion Island. Although being 40% (Table 2). Watson listed 20 endemic insects of of volcanic origin, rather than raised seafloor as is which a number have now been recorded elsewhere Macquarie Island (Williamson J 988), it is of similar or have been synonymised with other species. size, lies at a similar latitude Cfable 3), is of a Gressitt (1970) stated there were 2S endemic species similar age (Bakker et al. 1971) and the invertebrate of insects oul of a total of 40 (62%) including fauna is becoming well known (Crafford 1984, Collembola. There now appear to be only ten current 1986, Crafford & Scholtz 1987, Crafford et al. endemics (c. 18% - two Collembola, six Diptera Free-living terrestrial invertebrate fauna of Macquarie Island 37
TABLE 1 Terrestrial Invertebrates of Macquarie Island Excluding Mites
Order Number of species Number of species Number of native Total probably introduced introduced but and endemic number and now naturalised not naturalised species of species
Oligochaeta 7 8 Mollusca 1 3 4 Collembola 5 5 21 31 Psocoptera 1 Thysanoptera 1 Hemiptera 3 3 Diptera 2 12 15 Coleoptera 8 8 Lepidoptera 1 Hymenoptera 2 2 Araenae 2 2
12 6 58 76 + 88 mites
TABLE 2 Numbers and Percentages of Endemic Species on Macquarie Island*
Number of endemic species Percentage of endemic species
1967 & 1970 1987 1967 & 1970 1987
Oligochaeta 1 2 17 33 Mollusca 1 0 25 0 Collembola 3(9) 2+27t 33 6-12t Diptera 10 6 66 31t Thysanoptera 0 100 0 Psocoptera 1 0 100 0 Hemiptera 0 0 0 0 Coleoptera 0 12 Ot Hymenoptera 1+17 0 100 0 Lepidoptera 1 0 100 0 Araneae 0 0 0 Mollusca 0 0 0
Endemic species 20+17 10+27 -40% -17%
Total species 51 76
* Recorded by Watson (1967) and Gressitt (1970), and those currently known. t Taxonomic studies not yet complete. TABLE ., of Physical Characteristics of Marion and
Marion Island rie hland
2YU Penndl1cf:n iCe cover (ha) SO Area Gc:;iuw 'iOG nI (ha) 1SU 46.') of the (ulde-;( m,mlh i. DC) Distance from nearest mainland (km) lYO() 990 Di;"!:lfi.Ce froco nearesl land (kin) 1900 6iG DishlHce froln nearest land \vest\vard ! 0" latitude (km) :\270 MealJ a.I1Dual temperalure (OC) .'i. 1 tvlean annual (nun) 2575 Age million BP iniUion in' NIlInbtr of of v,u.;cular 22
'" Fwm Abboa (1974).
TABLE 4 of Number of insect Known from the Prine<: Edward and Island
Prim:;:: Edward hiland
Native and % endemic Naturalised Native am! 'Ii) el1demk Naturalised endemic aliens endemic aliens species (nos) species (nos)
(' o o ,J () () o o o 3 () 3 7 43(3) () o 5 2) 4 2 1 31( i) o o o o () 9 26
10
includes Marion Island Frum Crafford et ill. (ll)86) Free-living terrestrial invertebrate fauna of Macquarie Island 39
TABLE 5 Comparison of Collembolan Faunas of Subantarctic Islands
Island Total number Number of % endemic Number in common % of native and endemic species species with Macquarie Island endemic species
Macquarie 21 2(+2)* 10(720) Heard 8 1 12 7 88 Marion 13 2 13 7(+2)* 54 Kerguelen 30 9(+2)* 30 10(+2)* 33 Crozet 28 8 29 6(+3)* 21
* Species of Sminthurunus and Megalothorax not yet fully determined.
TABLE 6 Biogeographic Affinities of the Macquarie Island Terrestrial Invertebrate Fauna
Taxon Number Cosmopolitan Western New Zealand Unknown of subantarctic subantarctic species islands islands
Oligochaeta 8 1 5 2 Mollusca 4 1 1 2 Collembola 31 10 16 4 1 Insecta Psocoptera 1 Thysanoptera 1 Heteroptera 3 3 Diptera 15 3 2 5 5 Hymenoptera 2 1 1 Coleoptera 8 5 3 Lepidoptera 1 1 Araneae 2 1
Totals 76 18 24 20 14
and two Oligochaeta) with a few others doubtfully islands represent the summits of two extinct endemic (e.g. other Diptera). The Macquarie Island volcanoes and are separated by only 20 km. population of Eudonia (Lepidoptera) is distinct Although Prince Edward Island is only one-seventh probably at less than subspecific level (J. Dugdale, the size of Marion Island, they have practically pers. comm.). identical faunas. Crafford gives the percentage of This level of endemism is lower than that of insect endemics (excluding Collembola) for Marion Island and the neighbouring Prince Edward Kerguelen, Heard and Crozet Islands of 27, 11 and Island for which six (35%) of the 17 native insect 67 respectively. Only that of Heard Island, with a species, excluding Collembola, are endemic (Table small fauna of nine species and one endemic, is less 4). The other nine species are classed as naturalised than that of Macquarie Island. aliens by Crafford et al. (1986). Treated separately, As far as collembolan faunas are concerned, Marion Island and Prince Edward Island each have Macquarie Island has a similar level of endemism to only one species not occurring on the other. The two Marion Island, which is certainly under-collected, 4() Penelope Greenslade but less than half that of and Crozc, subantarctic islands to the west (V Island, (table 5). On rhe whole the Collembola can (Pseudosorensia) atlantica). In table 'i the relation· be taken as characteristic of the Illvertebratc fauna of of Macquarie Island Cnllembo!a with those of Macquarie Island because they are well other -.:ubantarctic islands is examined in n~OH~ knOVdl and collected and recent revisions arc delail. The Island fauna appears mo';[ availabic for the faunas of the Kerguelen group and Island, but thi" probabJ.v Heald hhnct, as \.yt'll as hiand. Also because the small fauna of the latter consis!~~ infomntion is availahle for Soulh aln)ost of a, Campbell and Auckland Islands. Therefore, estimates of endemism and should be reliable for this DISCUSSiON group. in the past, endemism has bet'n caiculated from knowll taxa, which can lead to bias When the Island fauna with (see Watson 1967, Gressit i Wallwork 1973, the fauna of thc other subantarctic islands, the many Abbott 1974). Collembo!a represent a high variables such latitude, altitude, proportion (40'10) of rbe total number of distance from land masses, climate, ice free molluscs and oligochaetes mites, from areas etc., have to he considered, Certain common Macquarie Island, The [rend in successive calcula features of their faunas do emerge in of those tions of the level of endemism has been lowards differences and these have been noted by earlier lowering this figure as faunas become better known authors (Gressitt 1961, 1962, 1964, 1970, Crafford taxonomically (table 2). et ai. 1986). An subantarctic islands have small faunas with a disharmonic of higher taxa 1974). The same characteristIcs have Affinities been noted for antarctic faunas (Block 1 However, the level of endemism varies, and on The taxonomic of the fauna are Macquarie Island, from which one endemic in table 6 where are as either plant is k!loWll (P. Selkirk, pe,s, comm.), the level of cosmopol ilan or eastern, western or endemism appears to be lower than might be unknown affinities. The classification is based on that the island is is well where the species or its closest relative occurs vegetated and has an equable climate .. Taking into out;,ide Macquarie Island, From the totals in table 6, account the trend towards a lower level of endemism it appears that the faunal affinities and hence origins with of the taxonomic lie approximately with the western and with affinities of the fauna, it is that it may fall the New Zealand subantarctic islands. As far as the even further. One explanation suggests that the insects afe concerned, relationships are largely with ecosystems on the jsland may be young (Selkirk et New Zealand. However, the Collembola and the al. 1983, Selkirk et ai. 1988), b(~en formed Oligochaeta show affinities to the west, i.e. with on a raised part of oceanic crust, most of which subantarctic islands on the SIde of the emerged above sea level the antarctic continent. These taxa are more Pleistocene (Williamson 1988), rather than being a associated with the soil than the insects which live relict of an older, once larger land mass. Gressilt for the most part above the soil and litter surface, earl ier recent origin for the for at least part of their life and tend to be Island fauna based on ihe few more that OCCllr there, From tablc (] it can be seen thal fOllr coli em- Island itself bad been part of a Dolan have affinities with New Zealand. mass (Gressitt 1(62). c;urface., leaf litter or above The lack of correlation in the Subantarctic (Lm(J\vsoni, I.lurhotti. P.davidi. between islanu area and number of ha~ been K.hattz
variable extent and it & Scholtz 1986) that preadapted for subantarctic habitats, Partheno can also be selected for in habitiats in which The [Sland fauna which ,ue been :j: in check Ii st in the appendix), Some of these parthenogenetic, for instance ali the and Chirotlomidae, Edwards & Usher (1985) have out that antarctic species have a need for flexible life histories, a characteristic of A selection. The composition of the fauna of Island is unlike any other subantarctic island in the lack of as an cllfculionids as noted by Gressitt ( 970), For the island have aIrnost instance, on all other subantarctic islands except recently introduced South Georgia several of curculionids occur become and they are often endemic. Three of the five curculionids in the Marion group of isiands are endemic and one of four on Heard Island terrestrial and marine ecosystems (Kuschel 1970). A suitable curcuhonid hahitat of marine invertebrates) ;\s a result 0 f in the P~Q roots in peat (a hahital ill the Falkland present. Islands described Lewis Smith & Prince (1985) as a "rernarkaole" environment) is present in abundance on Island. Possible reasons EPILOGUE for their absence are the cGmbination of poor dispersal ability and the island's "The ward, room IS tbe battle field of more scientific Alternatively, Crafford & SchGltz (19g7) thrills. The that the absence of a Shaddef(;n, Jock \V{)rciic. Robbie C~lark~ lIussey~ l)[ on Prince Edward Island and the Macklin and or two BlTned be the result of introduced mice .. Mice and as well as the insectivorous weka have bee a present on Island [9th century and It is po~sible they have had some effect on the in:\ect fauna but they arc of any The diet of Mus musculus consists mainly of IGSects i 986); arachnids, larvae and have been fOllnd in mouse stomach,; in that order of abundance, However, even where curculionids are abundanl subantarctic islands are not eJten any great extent by mice (Gleeson & van 1982) as the larvae of these insects live in soil and Free-ltl'ing terrestrial invcrtehratejiuma o/M{1cquarie Island 43
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EDWARDS, J.S., 1986: Derelicts of dispersal: arthropod fall Introduction. Pacif. Insects 3(4): 905-915. out on Pacific North West volcanoes. In GRESSITT, J.L., 1964: Insects of Campbell Island. Paci! Danthanarayan, W. (Ed.): INSECT FLIGHT: Insects Monogr. 7: 1-663. DISPERSAL AND MIGRATION. Springer GRESSITT, J.L., 1970: Subantarctic entomology, particu Verlag, Berlin: 196-203. larly of South Georgia and Heard Island. Paci! EDWARDS, M. & USHER, M.B., 1985: The winged Antarctic Insects Monogr. 23: i-iv, 1-374. midge Parachius steinenii (Gerke) (Diptera: HAMOND, R, 1987: Non-marine harpacticoid copepods of Chironomidae) in the South Shetland Islands. Australia. 1. Canthocamptidae of the genus Bioi. 1. Linn. Soc. 26: 83-93. Canthocamptus Westwood S. lat. and Fibula EVENHUIS, N.L., 1989: Appendix 1. Diptera of Antarctic camptus gen. nov., and including the description and Subantarctic Islands. In Evenhuis, N.L. 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HURLEY, F., 1979: SHACKLETON'S ARGONAUTS. THE Desidae, Dictynidae, Hahniidae, Amauro EPIC TALE OF SHACKLETON'S VOYAGE TO biodidae, Nicodamidae. Otago Mus. Bull. 3: ANTARCTICA IN 1915. McGraw-Hill Book Co., 1-184. Sydney. FORSTER, RR. & BLEST, A.D., 1979: The spiders of New JAMIESON, B.G.M., 1968: Macquaridrilus: a new genus of Zealand. Part V. Lynyphiidae - Mynogleninae. Tubificidae (Oligochaeta) from Macquarie Island. Otago Mus. Bull. 5: 1-173. Univ. Qld Pap. 3(5): 55-69. GLEESON, J.P. & VAN RENSBURG, P.J.J., 1982: Feeding JEANNEL, R, 1940: III Coleopteres. In Jeannel, R (Ed.): ecology of the house mouse Mus musculus on Croisiere du Bougainville aux lies Australes Marion Island. S. Afr. 1. Antarkt. Res. 12: 34-39: Fran~aises. Mem. Mus. Hist. nat. 14: 63-201. GREENSLADE, P., 1985: The Conservation Status of KUSCHEL, G., 1970: Coleoptera: Curculionidae of Heard Tasmanian Non-Marine Invertebrates. Unpub!. Island. Paci! Insects Mon. 23: 255-260. rep. Tasm. Nat. Pks Wild!. Servo LAWRENCE, R.F., 1971: Araneida. In Bakker, E.M. van Z., GREENSLADE, P., 1986: Additions to the collembolan fauna Sr., Winterbottom, J.M. & Dyer, RA. (Eds): of Heard Island. Rec. S. Aust. Mus. 19: 91-96. MARION AND PRINCE EDWARD ISLANDS. GREENSLADE, P., 1987a: Zoogeography of Tasmanian A.A. Balkema, Capetown: 301-313. COllembola genera. In SOIL FAUNA AND SOIL LEE, D.C. & HUNTER, P.E., 1974: Arthropoda of the Free-living terrestrial invertebrate fauna of Macquarie Island 45
subantarctic islands of New Zealand. 6. Rhoda SELKIRK, D.R., SELKIRK, P.M. & SEPPELT, R.D., 1986: An caridae (Acari: Mesostigmata). N.Z. 1. Zool. 1(3): annotated bibliography of Macquarie Island. 295-328. ANARE Res. Notes 38: 1-134. LEE, K.E., 1959: The earthworm fauna of New Zealand. SELKIRK, D.R., SELKIRK, P.M., BERGSTROM, D. & NZ. DSIR Bull. 130: 1-486. ADAMSON, D.A., 1988: Ridge top peats and LEE, K.E., 1968: Oligochaeta from subantarctic islands. palacolace deposits on Macquarie Island. Pap. BANZARE Rep. Ser. B8(8): 149-165. Proc. R. Soc. Tasm. 122(1): 83-90. LEWIS SMITH, Rl. & PRINCE, P.A., 1985: The natural SELKIRK, P.M., SEPPELT, R.D. & SELKIRK, D.R., 1990: history of Beauchene Island. Bioi. 1. Linn. Soc. SUBANTARCTIC MACQUARIE ISLAND: 24: 233-283. ENVIRONMENT AND BIOLOGY. Cambridge MACARTHUR, R.H. & WILSON, E.O., 1967: THE THEORY University Press. OF ISLAND BIOGEOGRAPHY. Princeton SIMS, R.W., 1971: Oligochaeta. In Bakker, E.M., van Z., University Press, Princeton, N.J. Sr., Winterbottom, J.M. & Dyer, R.A. (Eds): MCQUILLAN, P.B. & MARKER, P., 1984: The kelpflies MARION AND PRINCE EDWARD ISLANDS. (Diptera: Coelopidae) of Macquarie Island. Tasm. A.A. Balkema, Capetown: 391-393. Nat. 79: 17-20. SMIRNOV, N.N. & TIMMS, B.V., 1983: A revision of the MATHIS, W.N., 1989: Coelopidae. In Evenhuis, N.L. (Ed.): Australian Cladocera (Crustacea). Rec. Aust. CATALOGUE OF OCEANIC AND AUSTRAL Mus., Supp. 1: 1-132. ASIAN DIPTERA. Bishop Museum Press, SMITHERS, C.N., 1964: Insects on Campbell Island. Honolulu and EJ. Brill, Leiden: 563-564. Psocoptera. Pacif. Insects Monogr. 7: 226-229. MATSUDA, R., 1976: MORPHOLOGY AND EVOLUTION SMITHERS, e.N., 1972: The classification and phylogeny of OF THE INSECT ABDOMEN WITH the Psocoptera. Mem. Aust. Mus. 14: 1-349. SPECIAL REFERENCE TO DEVELOPMENTAL STEEL, W.O., 1964: Insects of Campbell Island. PATTERNS AND THEIR BEARINGS UPON Coleoptera: Staphy1inidae. Pacif. Insects Monogr. SYSTEMATICS. Pergamon, Oxford: 1-534. 7: 340-375. MOUND, L.A. & WALKER, A.K., 1982: TEREBRANTIA STEEL, W.O., 1970: Coleoptera: Staphylinidae of South (INSECTA: THYSANOPTERA). FAUNA OF NEW Georgia. Pacif. Insects Monogr. 23: 240-242. ZEALAND 1. Wellington, New Zealand: 113 pp. SUBLETTE, J.E. & WIRTH, W.W., 1980: The Chironomidae NEWTON, A.F., 1985: South temperate Staphy1inoidea and Ceratopogonidae (Diptera) of New Zealand's (Coleoptera): their potential for biogeographic subantarctic islands. NZ. 1. Zool. 7: 299-378. analysis of austral disjunctions. In Ball, G.E. THORNTON, LW.B., 1985: The geographical and ecological (Ed.): TAXONOMY, PHYLOGENY AND distribution of arboreal Psocoptera. Ann. Rev. ZOOGEOGRAPHY OF BEETLES AND ANTS. Entomol.30: 175-196. Junk, Dordrecht: 180-220. WALLWORK, J.A., 1973: Zoogeography of some terrestrial PrERREHUMBERT, C.P.H., POWELL, F.A. & OLIVER, S., 1985: micro-arthropoda in Antarctica. Bioi. Rev. 48: Continental transport of particulate matter 233-259. between Australia and Macquarie Island. Proc. WASHBURN, J.O. & WASHBURN, L., 1984: Active aerial 8th Int. Clean Air Conf., Melbourne, 1984: dispersal of minute wingless arthropods: 741-750. exploitation of boundary layer velocity gradients. QUATE, L.W., 1962: Insects of Macquarie Island. Diptera: Science 233: 1088-1089. Psychodidae. Pacif. Insects 4(4): 958. WATSON, K.C., 1967: The terrestrial Arthropoda of RAINBOW, W.J., 1917: Arachnida from Macquarie Island. Macquarie Island. ANARE Sci. Rep. Ser. B(1),99: AAE Sci. Rep., Ser. C5(1): 1-13. 90. RAMAZZOTI, G. & MAUCCI, W., 1983: IL PHILUM WATT, J.e., 1971: Entomology of the Aucklands and other TARDIGRADA. III. Elizione Riveduta e islands south of New Zealand: Coleoptera: Aggiornata Memoire dell Instituto Italiano di Scarabaeidae, Byrrhidae, Ptinidae, Tenebrionidae. Idrobiologia, Verbania Pallanza, Italy: 1012 pp. Padf. Insects Monogr. 27: 193-224. SABROSKY, C.M., 1962: Insects of Macquarie Island. WIGGLESWORTH, V.B., 1952: Hormone balance and the Diptera: Chloropidae, Milichiidae. Pacif. Insects control of metamorphosis in Rhodnius prolixus 4(4): 973. (Hemiptera). 1. Expl. Bioi. 29: 620-631. SALMON, J.T., 1964: An index to the Collembola. Bull. R. WILLIAMS, R., 1988: The nearshore fishes of Macquarie Soc. NZ. 7: 651 pp. Island. Pap. Proc. R. Soc. Tasm. 122(1): 233-245. SEGUY, E., 1940: IV Dipteres. In Jeannel, R. (Ed.): WILLIAMSON, P.E., 1988: Origin, structural and tectonic Croisiere du Bougainville aux lIes Australes history of the Macquarie Island region. Pap. Proc. Franc;:aises. Mem. Mus. Hist. nat. 14: 203-267. R. Soc. Tasm. 122(1): 27-43. SELKIRK, D.R., SELKIRK, P.M. & GRIFFIN, K., 1983: WISE, K.A.J., 1977: A synonymic checklist of the Palynological evidence for Holocene environ Hexapoda of the New Zealand sub-region. The mental change and uplift on Wireless Hill, smaller orders. Bull. Auckl. Inst. Mus. 11: 1-176. Macquarie Island. Proc. Linn. Soc. NSW 107: WOMERSLEY, H., 1937a: Acarina. AAE Sci. Rep., Ser. 1-17. C10(6): 1-241. 46 Penelope Greenslade
WOMERSLEY, R, 1937b: Coleoptera. BANZARE Rep. Sa \7;/O]\1ERSL.EY, rI.. 1937d: Diptera. Bllj\/ZA/Qr Rep B4(l): 23-26. B4(3): 59-79.
WOMERSLEY j l-I., 1937c: Collembola. BA.NZAR/~' Rep., 5'pr. YUSHIMOTO, C.M .• 1962: Insi.~ct:.. of iVbcquaric rslarrd. B4(l): 1-7. Diapriidac. Poc/,f. insects 4(4)'
APPENDIX Annotated Checklist of Terrestrial Invertehrates of Isiand
* new record (localities given) ~ native (indigenous) t endemic species :l" introduced and natu raj i sed # species recorded earlier from Macquarie Island introduced and not naturalised but probably misidentified
OLIGOCHAETA NIOLLUSCA MEGASCOLECIDAE ATHORACOPliORIDAE Microscolex macquariensis (Beddard 1896) -r Refiectopallium marrensi (Suter 19(9) ~ Considered to be endemic to Macquaric Island (Lee 1959, Athoracophorus (Amphiconophora) Suter 1968) and, according to Lee, most closely related to species 1.909 from South Georgia and the Kerguelen Islands, but also Also occurs on Auckland Island. An indigenous specites close to a species in Campbell Island. Sims (1971) states according to Watson (1967) but not recorded by Dell that this species may only be a geographical race and not (1964). Burton (963) erected the genus Relleetopal/ium a fu 11 species. and gave the Auckland hlands as the locality but LUMBRICIDAE subsequently mentioncd a Macquarie Island iocality (p. Bimastus tenuis (Eisen 1874) 73). In a later publication (Burton 1980) he added a further Recorded by Lee (1968) and also known from Kerguelen discussion of the species. and Heard Islands; a cosmopolitan species. Pseudaneitea huttoni (Suter 19(9) 41 = TUBIFICIDAE Alhoracophorus (Pseuduncilea) huttoni Suter 1909 Macquaridrilus hennettae Jamieson 1968 Also occurs on The Snares. An indigenous species An endemic species of a monotypic genus. It is considered according to Watson (1967) but nOl recorded by Dc!! to be an ancestral form of obscure relationships and may (1964). Burton (196]) did not give a Macquarie Island have reached the island only recently (Jamieson 1968). locality. ENCHYTRAEIDAE LIMACIDAE Enchytraeus alhidus Henle 1837 ~ Deroceras reileu/aris MUller 1774 j: = Agrioiirrun Also known from the Kerguelen and Crozet Islands; agrestis Linne sensu Hedley 1916 recorded from Macqnarie fsland by l.ee (1968) {'rom a According to Hedley (1916), this European slug has been determination by L. Cernosvitov; occupying marine, introduced and was first found around scalers' huts at limnic and terrestrial habitats. One of the commonest Lusltania Bay. 11 is the cornmcncst British slug and has u species in the family. cosmopolitan distribution. Dell (1964) commcmed 1halit Pachydrilus macquariensis (Benham 1915) = is now well established. Lumhrieillus macquariensis Benham 1915 ~ PUNCTIDAE Also known from Heard Island. Recorded by Lee (1968) Phrixgl1athus hamilroni Suter I R961[ from a determination by L. Cenosvitov. Looma (Phri.x:r;nathus) harniltoni SUTer lx96 Pachydrilus maximus Michaelsen 1900 ~I Phrixgnalhus hand/rom Hedley 19167 = Also known from Heard Island and the Kerguelens. Laoma campheiiica Hamilton 1894 (mis Recorded by Lee (196R) from a determinmion by L. identification) Cernosvitov. According to Hedley (19 j 6), (his snarl was tbe Pachydrilus werthi (Michaelsen 1905) = Lumhricillas "southernmost in the world", but molluscs have now been werthi (Michaelsen 1905) II recorded from terrestrial habitats on Heard, Possession. tbe Marionina antipodum (Benham 19(5) ~ Kerguelen, Marion, and South Georgia (Cumberland Bay East) Islands (Block 1(84). Dell (1964) considered that this species is very close [0 an inadequately described specjes front Cmllpbell ]stand, Phn'xgnathus cam!7f'llic(l /~f IVI ,']{'(j1",mH' 1sf all~~l
Zealand. F. Chr(10 (per:--., comrn.) consider'; the CarnpbeiL Auckland .Nlw.:qu:1rle specirnens tu be although sp,_>.:.:ie~; rnorphoJogicrdl,y variable. IsofOrna (F'[J'J'Ortlotorno) Jlunctata Wahlgre n
TARDIGRA Dil, in lhe Not l)n A'u~:kldnd MACROBiOTlDAE Carnpbcl! ir.;iands, where a different Uf1,nnINU\ aUl:usti (l\;lunav 19(7) ~? occurs.
1(83). collected, as DHmy been publislied Neariy ,,,1 "pped[ is(){{J!na (D. Horning, comrn.L 1970) t
ARTHROPODA arc, required. n.'n/,,,,, "",,1\' antorclicus Willem n()1 COLLEMBOLA A C0J111110n species of the Antarctic and Subantarctic HYPOGASTRUR1D,,\E Cryp/opygl4s dunius Deharveng 1981 ~ Hypoga,vtrura (fJypogastFura) purpures'cen,)' Also known frorn Marion Island. (Lubbock 1868) j: Wahlgren 1900 ~ f/ypor;asrrura (HVl'ogasil'llra) vialiea (Tul1berg A very widespread ~pecies ;1Dd native to the 1872) l!emisphere, f/ypogastrurrl (Cerarophysel/a) dentieulata (Bagnall Cryptopygus' !awren(J!i Deharveng I 981 ~ 1941) Also knov;n Kergue]en IslanrL These three species arc cosmopolitan and 1-/ purpurescens Cr.-vpiOp}'gus tri('uspis Enderf,;:,in ] 909 ~ and H. denticrdata have probahly been n~cently Widespread in Subantarctic (Greenslade 19S6. (Greenslade Wise] 9x4. 198()1. NEANURlDilE Friesca Ii/hrooki Wise 1970 ~ 'viennei station greenhouse, out a COfYIn1(m Dcharveng 1981 nurncroutj records {roin Australia A1so known Heard Island and Bouvetoya (Greenslade 1986. Greens];,de & 1996) Is()/ SM1'JTHUR1DAL Rh()pal(}.)i[J/li,r!l Sfltinflwrides ,-,p but is rarc group OnJy i~nO\1'n group ('dIVer, f)t"r:." '0 1960, hCIXC: ju: Gtcenc;lade, in pH.:p.) only (1982) reci)nl C:HIll' be 1I Island, was collccrcJ. Zealand, on Ntacquaric Slfr inthurituls' kcri'J'u'if'n,'!1\11' COLEOPTERA: STAPHYLlNlDAE A 1so known frorn OM:ALHNAF iden:tfic~d from ~1acqtlari\~ Omaliomimu5 aihipelille (Kiescnwetter 1877) ~ an inltl1atufc spccirnen Omaiium aihipnlnP KicsenweUer 1877 = Omalium ioentifi"ation, l'ariipenllc Lea 1920 nec Womersley1937 = ?S'rninrhurinus cr. rerrr),\tri.'i' V~lorncr~~1ey ! 931 ~l?t Omaliomimus jlavipcnnis 1947 Collected sporadIcally Also known from Auckland and Campbelllsiunds (Jeanne! Snlilithurlnus rm{j{jniY1m ]940, Gressill 1970); wingless Only blown froID the Omaliomimus ,'enaior (Broun 19(9) 11 prevHJusly only recor(Jcd 1909 S'minthur inu,'i' (;p. cL & Aho known frorn and Campbell Islands (Steel Rapoport 1963 1964, \Va1son t9{,7, Gressit1 1970); a wingless species. Also known lr0111 hern South Arrlcrica. V:H10H:l SZfllomaliu!11 (Camero!! 1945) II collection;;.: from the Plateaa and fronl coast<-li sites. Onwlium helms! Cameron 1945 sp. granulosus Endcrlcill l009 Als0 known from Zealand (Watson 1967, Gressitr Also recorded frum JCcrguclen <.ind ~1anon 1970); winged "nd often found on carrion (M, Thayer, lslandc; (De Harveng 19K1), Only specimens arc comrn., Newton 1985 Ornaliinae Genus E). avallah]I:; so a definite identification cannot he made. Stenoll1aiium suicithorax (Broun l880)~ Locality: POll ~Jn Ha~"selborcugh Bay Omafium ,)'u/CitJ'o!'ax Broun 18HO ;::- Omaliwrt 250 lll. perp!cxum Broun 1xQ4::: flomat'ium varl;pe1me sensu Kat/anna harL-anJ SdlmoH 1\)64 t Womerslcy 1937 nee Leo 1920 At present this spec]e~ only known frorn Iv1at'quarie Watson (1967), in publishing Ibis synonymy, quoted W,O hland (rod lTIay e!ldernic, although unidentified Steel's unpublished opiniom thar the specimens studied by immature Kalianuo ;~pecics h~\vc bel,~n fnJ1.11 Catnpbcll Womcfsky (19371;;) and identified by him as A. Island and ~lre kno\"'rl frorn ihe Kerguelen-; ;lnd The Lfozcntpnsis Enderkin were neither that species nor NELLlUAE Lea baT were conspccific with 0. perplexurn A1eguio/hol'{lX sr. Brenm. The specje:~ !:; winged and known froTn carrion. Not iclenti fied a, The; kJIOWn worldwide. Also found in New leu land. Crymu,\ ::-...:" Afpediomirruf,,,' sp, INSECTA Recorded by Skf'l (1 Y64 :149, Nt Thayer, pel's. COrI1ITJ.), Pson)PTERA: wJngs vestigiaL /:.ustropsocus in.1H/{;ri,:y SmilheL'; l!{}riipen!1(; Til1yard 1Q20 j nee L.ea 1920 Recol ded by '-''larson (1967) Probably larva or identificution (M. Thayer, frOID n~G Antipodes, CanlpbeH :Jnd Auckt8W.:l comm.L ISland.., (Srnilben; 1964, \Vl~',e 197/). The :~evenll ALEOCHARIN AE specie~j in Nevi ~.tt't1jar:d (Thornton 1985), fialmapusa anlarctif'(j ICeSeuwcttcr i 877 :::: HEM1PTERi\: APWDiDAE Antarctophytosus macquariensis Womerslcy 1937~ }ucksonia rJ(lpiflala Theubald 1923 Also knovJI1 from Auckland Island (Steel 1964, J no, This ~clpeclcs occurs EUHipe and ['.Jew Zealand (Eas1op 'Watson J 967), The species is wingless and terrestrial. 1962, ! 970), bur apparently absent from Auslra!ia, In Europe it appears tD prefer cooL hmnjd (IvL The Staphylinidae (OmaJiinae) are currently being Carver) pers. corntn.). studierl hy 1\1, T11"yer, Free-living terrestrial invertebrate fauna of Macquarie Island 49 BYRRHIDAE species is winged while Heard Island has wingless endemic Epichorius sorenseni (Brookes 1951) ~ in this family. Only one adult, one pupa and some larvae known (Watson 1967), identified by Watt (1971). Also known from Evenhuis (1989) records another species from Campbell Island. Macquarie Island, Sciara womersleyi Seguy 1940. This DIPTERA: TIPULIDAE species was erected by Seguy for an unnamed sciarid Erioptera (Trimicra) pilipes macquariensis described by Womersley (l937d) from two females from Alexander 1962 = Trimicra pilipes Fabricius 1787 ~ Kerguelen Island. Womersley also had a single sciarid The nominate species is cosmopolitan. This may be a from Macquarie Island which he believed was the same variety rather than a subspecies as interpreted from species. Because S. womersleyi was inadequately Watson (1967) and P. Johns (pers. comm.). The species is described, it must remain species inquirenda for the known to be variable (P. Johns, pers. comm.), and present. (D. Colless, pers. comm.) Macquarie Island specimens exhibit much morphological variation (Alexander 1962) which is within that of DOLICHOPODIDAE Campbell Island and New Zealand populations (P. Johm, Schoenophilus pedestris pedestris Lamb 1909 t pers. comm.). A winged species. Redescribed by Harrison (1959) and endemic according to PSYCHODIDAE Harrison (1976). Another subspecies occurs on Auckland Psychoda surcoufi Tonnoir 1922 = Psychoda and Campbell Islands. Wings reduced. spatulata Satchell 1950 = Psychoda subimmaculata HELCOMYZIDAE Tonn. sensu Womersley 1937 :j: Paractora asymmetrica (Enderlein 1930) ? = This species is also known from New Zealand (including Actoceles asymmetrica Enderlein 1930 Campbell, Auckland and the Antipodes Islands), southern Not seen since the original collection which has not been Australia and southern South America (Duckhouse 1985). restudied. There is doubt as to the family placing of this Psychoda parthenogenetica Tonnoir 1940 :j: species (Harrison 1976). Psychoda severini parthenogenetica Tonnoir 1940 COELOPIDAE A species restricted to Europe, Africa, Japan, Australia and Coelopella plebeia Malloch 1933 ~ = Coelopa New Zealand (Duckhouse 1970,1971,1985, Watson 1967) macquariensis Womersley 1937 and known also from Crozet, Marion, South Georgia and This species is restricted to Macquarie Island according to Kerguelen Islands. Mathis (1989) who is not in agreement with Harrison's Psychoda alternata Say 1824 § (1976) synonymy of this species with Coelopa Fucomyia There is only a single record of this cosmopolitan species curvipes Hutton 1902 from Auckland and Chatham Islands from Macquarie Island (Quate 1962) and this species is and New Zealand. McAlpine currently revising the probably not a permanent resident of the island Australian Coelopidae. (Duckhouse 1985). Icaridion de bile Lamb 1909 ~ = Coelopa nigrifrons CHIRONOMIDAE Lamb 1909 Telmatogeton macquariensis (Brundin 1962) = According to Harrison (1976), this species occurs on Helirytus macquariensis Brundin 1962 t Campbell and Auckland Islands, the Antipodes, The An endemic species according to Watson (1967). Brundin Snares and mainland New Zealand as well as Macquarie (1962) said it is closely related to the Kerguelen species Island. McQuillan & Marker (1984) omitted to record the Telmatogeton amphibius (Eaton) with which Sublette & synonymy made by Harrison (1976). Wirth (1980) agreed and mentioned that it is also close to CARNIDAE a species from the Antipodes Islands. Wings strongly Australimyza macquariensis (Womersley 1937) t = reduced. Procenace macquariensis Womersley 1937 Smittia sp. ? Endemic according to Harrison (1976); it is common and Recorded by Brundin (1962) who believed it to be belongs to an endemic genus for the New Zealand bio restricted to Macquarie Island; possibly a parthenogenetic geographic region. Another species in the genus is found in species that has bever been described. (Specimens labelled Antipodes, Auckland and Campbell Islands and The Pseudosmittia sp.). Snares. EPHYDRIDAE Two further species of chironomid were collected in the Ephydrella macquariensis (Womersley 1937) t = summer of 1986-87 in two different genera and are Ephydra macquariensis Womersley 1937 currently being studied by P. Cranston. An endemic species for Macquarie Island according to Harrison (1976); Bock (1987) recorded it erroneously from SCIARIDAE New Zealand subantarctic islands. Bradysia watsoni Colless 1962 t Amalopteryx maritima Eaton 1875 # An endemic species according to Watson (1967) but there This species is also known from Heard, Crozet and the are many undescribed species of sciarids in collections Kerguelen Islands according to Womersley (1937d) but is from neighbouring regions (D. Colless, pers. comm.). This a misidentification for Apetaenus watsoni according to 50 \ 14(2) TET1HNHlAF Apett:{(!I1US H-'{?t.voni A \vingk:~~s specie;;; and Island ccording i,O but j"he record is l!sLCd litorali"\', CHLOHOPTDAE ThyriduJa sp, Recorded by Sabrosky and known fr01l1 only \,/,;0 \VeTe not by Bani';eD (1976), LEPIDOPTERA: PYRALJDAE LYNPI1HDAf: Eudonia rtl{J1vsoni (\V()mersky 1(37) Scoparia rnawsoni Wornersley & Tinda1e 1937 nov. cornb. Also Auckland, /\.nUpodes and Redescribed by COlTlfll0n (1962) and considered endenlic, Enderby Island:; (Hic.kman [939, VVatson 1967, Forster & althougb J Dugd;:tle COITlIn.) has cOHlpared Blesi 197'1) specimens with congenerics from Zealand alld its flaplinus ru/[)(ephaiia (Urquart 18SB) # subantarctic is1ands and states MYl10gienes (Urquarl j 8S8) ~ readily distinguished Al,vtloglenes insolefIs Simon 1905 psammitis fratn SOllth Island". Macqunrie Recorded HickrnaIl (1939) and fron") spt:cimcns belong to the New Zealand wide L, Allele bnd (Meyrick) group of popuiation-; this "species has be(;Il In!sidcntifie·d, tendency to have l()(.:al populutions" (T. Dugdale, in litL), The Macquarie Island populuioll is therefore more sp. #'1 accurately described as a geographical race rather than ;3 Forster (I')S~) recorded a Hahnia Frorn _Auckland distincl species, Generic name lIlcorrectly used (l (31)" recorded Common (1962) Jnd Watson (1967) (M. frorn Macquaric No trace of record can Sl1afrec in btL). bc found and conc1udc that Forster mistakenly i-lickrnanls (l9~j9) des<;riptions IfaJ/nia HYMENOPTERA: DIAPRIIDAE frcnn Crozet Island. Antarctopria latig(lster Brut's i 920 ~ Recorded hy Yoshimoto (1962) and redescribed by Early two of spidcr~, Par(41~oneta tnorrincri 8nd (1978, 1.980), AJ so known froIll Stewart, Antipodes, l!aplinus n4()cephalia were coilected in 1986;R7. Both Auckland and CampbelJ Islands, and the NevI were comn10n and abundant ZealfEld inainland.