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Memoirs of the Museum of Victoria 56(2):557-56l (1997) 28 February 1997 https://doi.org/10.24199/j.mmv.1997.56.50 DIVERSITY AND BIOGEOGRAPHY OF AUSTRALIAN (DIPLOPODA)

Dennis G. Black

School of Zoology, La Trobe University, Bundoora, Vic. 3083. Australia

Abstract

Black, D.G., 1997. Diversity and biogeography of Australian millipedes (Diplopoda). Memoirs of the Museum of Victoria 56(2): 557-561. Twenty families of millipedes, representing nine orders, have been recorded from Aus- tralia. Three of these families and one of the orders are introduced. The greatest diversity is found east of the Great Dividing Range and in the south-east, including Tasmania. Approxi- mately 250 native species have been described, probably representing only 10-20% of the actual number. Large areas of Australia, such as the north-west and west central coast, have pre- yet to be adequately surveyed. Lack of detailed taxonomic and biological information cludes use of Australia's diverse fauna for environmental management pur- poses.

Introduction taxonomists working within Australia. All taxo- nomic descriptions of Australian millipedes of Australia is rich but The millipede fauna published before the 1990s had been prepared below, nine of the largely unknown. As detailed by European and North American specialists. the world fauna 15 orders recognised in Many of the specimens on which these works of the approximately (Hoffman, 1979) and 20 were based were collected during major expedi- 1982) have been 100 families (Hoffman, tions to Australia early this century (Atterns, Three of the 20 famil- recorded from Australia. 1911; Verhoeff, 1924). No attempt had been orders have been intro- ies and one of the nine made, however, to characterise the nature of the settlement. P.M. Johns duced since European fauna as a whole. Modern millipede studies estimates that there are at least (pers. comm.) began after the visit in 1 980 of C.A. W. Jeekel, an times the pres- 2000 native species, about eight eminent European systematist. In reporting this majority of the undes- ent known fauna. The visit, Jeekel (1981) summarised the state of taxo- soil and cribed species are small, inconspicuous nomic knowledge of eastern Australian milli- litter dwellers. pedes and documented zoogeographic distri- work were fourfold: The aims of the present butions of higher taxa. His summary is, and will the history of Australian 1 to briefly summarise continue to be an invaluable reference for all millipede studies; subsequent millipede studies in this country. overview of the taxonomic and 2. to give an In the 1 980s, most published accounts dealing Australian milli- morphological diversity of with Australian Diplopoda by local researchers pedes at the order and family levels; concerned the ecology and control of the intro- geographical distributions 3. to document the duced Ommatoiulus moreletii Lucas, 1 860 (the regionally of millipede orders and families Black Portuguese Millipede). The research was within Australia; and concentrated in South Australia, where localised in our present 4. to identify the major gaps outbreaks of this nuisance pest were considered Australian millipede knowledge of the serious enough to have warranted considerable fauna. efforts in a search for a biological control agent. based on data Distributional records are The search was carried out both in the area of lodged in all of gleaned from specimen material origin of the species (Iberia) and in south-eastern supplemented the major museums in Australia, Australia, and resulted in the importation of a collections made by the with records from parasitic fly from Portugal and the identification research collections of author and the personal of a nematode parasitic in Australian millipedes Majer in R. Mesibov in Tasmania and J. (Baker, 1985a, b; Bailey, 1989; Shulte, 1989). Western Australia. Possible interactions between O. moreletii and native millipede species were investigated as an Australia Millipede studies in honours project at Flinders University in millipede Adelaide (Griffin and Bull, 1995). Prior to about 1990 there were no

557 558 D.G. BLACK

Other applied work on millipedes in Australia key for identification of millipede material to has dealt with the ecology of the pincushion mil- order level. lipede Unixenus mjobergi (Verhocff, 1924) in The pincushion millipedes (Polyxenida) are the Pilbara region of Western Australia. Like O. atypical, being very small (generally less than moreletii this species also becomes a nuisance 5 mm in length), extremely setose and able to when it invades townships in the hundreds of withstand much drier conditions than most thousands after rain (Burt, 1984; Koch, 1985). other Diplopoda. Australian species of the order Chordeuma- Pill Millipedes () are inhabi- tida (Craspedosomatida) have received recent tants of wet eucalypt forest and rainforest. Since attention (Golovatch, 1986; Mauries, 1987; rainforest invertebrates have been much better Shear and Mesibov, 1994, 1995), after first collected in Australia than those from many having been mentioned as being present in Aus- other habitat types, the systematics of the Aus- tralia by Jeekel (1981). This order is well rep- tralian pill millipedes is probably better known resented in Australia, but all species are cryptic than that of any other order. litter dwellers. The fact that it is only now being The orders Polyzoniida and Siphonophorida well documented illustrates the embryonic state are known as 'sucking millipedes' and show a of our knowledge of Australian millipedes reduction in mouthparts which is much more generally. pronounced in the latter. Both groups are pri- Studies by resident Australian taxonomists marily wet forest millipedes. They are obviously have only begun in the last five years. R. not typical macrodetritivores, given the struc- Mesibov (Queen Victoria Museum and Art Gal- ture of their mouthparts, but what they do eat is lery) has contributed to works on the Tasmanian unknown. Chordeumatida (Shear and Mesibov, 1994, The Chordeumatida are small litter dwellers 1 995) and is investigating the systematics of the found in wet forest. Members of the two families Tasmanian Dalodesmidae (Polydesmida) and found in Australia can be distinguished from the zoogeography of all Tasmanian millipedes. other millipedes by the occurrence on each body The author has undertaken revisions of genera segment of six dorsal macrosetae. in two Australian families, Siphonotidae (Poly- The most taxonomically diverse of the milli- zoniida) and Siphonophoridae (Siphonopho- pede orders in Australia is Polydesmida. The rida). and is conducting a family level review Paradoxosomatidae is the most speciose of all and producing a key to the millipede families the native millipede families and paradoxoso- occurring in Australia. matids are common in most forest and wood- Recent investigations of cave millipedes at land habitats. Dalodesmidae are common in the Cape Range and Barrow Island, Western Aus- east and south-west, and are particularly well tralia, were initialed by W. Humphreys at the represented in Tasmania. Members Of the Western Australian Museum (Humphreys and families Haplodesmidaeand Pyrgodesmidaeare Shear, 1993; Hoffman, 1994). Allozyme electro- difficult to distinguish from one another. They phoresis carried out on cave populations of the arc small, highly sculptured and rigid-bodied polydesmidan Stygiochiropus spp. from Cape , widespread in soil and litter. Very little Range (Humphreys and Shear, 1993) is one of is known about them in Australia, and there may the few examples of a molecular approach to be many undescribed species. At least one millipede systematics in the literature. species of the family Polydesmidae, Brachycies-

mus superus Latzel, 1 884 has been introduced to Tasmania (R. Mesibov, pers. comm.). The I axonomic and morphological diversity occurrence of this or other introduced Polydes- Orders and families of Australian millipedes are mida elsewhere in Australia has not been listed 1 in Table . Omitted from this table are the reported. introduced taxa including the order Julida Along with the introduced Julida, the last two (families Julidae and Blaniulidae) the and Poly- orders in Table I are collectively known as desmidae of the order Polydesmida, leaving 'juliform', which refers to the general body eight orders I and 7 families. Only one family, shape. Australian species of Julida are intro- Peterjohnsiidae, is known to be endemic (Mau- duced. Most are restricted to metropolitan areas ries, 1987). The family list in Table 1 is based in southern Australia, but Ommatoiulus morele- primarily on Jeekel 1 ( 98 1 ), Hoffman ( 1 982) and tii has made substantial inroads into dry native the unpublished work of P.M. Johns. Harvey bush around the cities. The three families of and Yen ( 1 provide 989) a very useful illustrated in Australia are typically found in DIVERSITY AND BIOGEOGRAPHY OF AUSTRALIAN MILLIPEDES 559

Table 1. Regional distributions of Australian millipede families, excluding those introduced. P = present (recorded from that region); Qld = Queensland, NSW/Vic. = New South Wales and = Victoria, Tas = Tasmania, NT = Northern Territory, SA = South Australia, Upper WA upper Western Australia, Lower WA = lower Western Australia. Boundaries arc approximate, based on divisions shown in Figure 2.

Taxon Qld NSW/ Tas. NT SA Upper Lower (Orders in capts) Vic. WA WA POLYXENIDA Synxenidae P P P P P Polyxenidae P P P P P P P Lophoproctidae P P SPHAEROTHERIIDA Sphaerotheriidae P P P P SIPHONOPHORIDA Siphonophoridae P P P POLYZONIIDA Siphonotidae P P P P P P CHORDEUMATIDA Peterjohnsiidae P P P Metopidiotrichidae P P P P POLYDESMIDA Paradoxosomatidae P P P P P P P Dalodesmidae P P P P Haplodesmidae and Pyrgodesmidae P P P P P P SPIROSTREPTIDA Cambalidae P P P Iulomorphidae P P P P P P P SPIROBOLIDA Rhinocricidae P P Pachybolidae P P P P P Spirobolellidae P P

of difficulties in assign- rotten logs or litter of wet forest in the north of for this analysis because the other, giving the continent. The two families of Spirostrep- ing species to one family or of eight orders and tida are widespread, with many species adapted maximum possible numbers families. to drier conditions than are suitable for other 16 survey work is carried millipedes. They are conspicuous components As more collecting and remote areas of Australia, the num- of the forest floor fauna, being found commonly out in more

bers in Table 1 are likely to increase consider- in litter, rotten wood and under bark at the bases ably for the central and north-western divisions. of trees. However, our present knowledge of distri- the greatest millipede Distributions of orders and families butions suggests that diversity occurs in eastern Australia. This domi- the orders and 1 lists the distributions of Table nance is almost certainly associated with the Australia within families of millipedes found in widespread occurrence of wet forest types east of of the continent (Tas- six arbitrary divisions the Great Dividing Range. mania constituting a seventh region). The three been omitted from the introduced families have Gaps in knowledge compilation. The table provides an overview of There is still a great deal of taxonomic work to be the diversity of Diplopoda at higher taxonomic done on Australian millipedes. Polyxenida are levels in different areas of the country. Pyrgod- widespread, but seldom collected esmidae and Haplodesmidae have been lumped common and 560 D.G. BLACK

or recognised as millipedes. Large collections of to me, to the Australian Biological Resources material do exist, however, and the handful of Study for providing support for field work and species now known will no doubt increase to museum visits and to R. Mesibov and J. Majer many times that number when Polyxenida for allowing me access to their research collec- eventually receive some attention. Sphaerother- tions. R. Mesibov also provided many helpful iida are reasonably well known, and revisions of suggestions as a referee of the manuscript. the Polyzoniida and Siphonophorida are being completed. Chordeumatida are currently receiv- ing attention, but there are still many species to be described. Within the Polydesmida, the References Dalodesmidae are currently being looked at, but the Paradoxosomatidae, the largest of all Aus- Attems, C, 1911. Ergebnisse der Hamburger Sudwes- tralian families, are not being studied. The two taustralischen Forschungreise 1905. Myriopoda. families of tiny Polydesmida, Pyrgodesmidae Die Fauna Sudwestaustralischem 3(6): 145-204. Bailey. and Haplodesmidae, are essentially unknown, P.T., 1989. The millipede parasitoid Pelidnop- tera nighpennis (F.) (Diptera: Sciomyzidae) for although they are clearly well represented in the biological control of the millipede Ommatoiu- Australia. Both orders of native juliforms, lus moreletii (Lucas) (Diplopoda: Julida: Julidae) Spirobolida and Spirostreptida, are badly in in Australia. Bulletin of Entomological Research need of work, with no resident expertise avail- 79: 381-391. able. Baker, G.H., 1985a. Parasites of the millipede Omma- As limited as our taxonomic knowledge is, our toiulus moreletii (Lucas) (Diplopoda: Iulidae) in knowledge of the general biology of millipedes in Portugal, and their potential as biological control Australia is even more sparse. Most of what little agents in Australia. Australian Journal ofZoology 32: 811-822. we do know was collected in the course of Baker, G.H., 1985b. The distribution and abundance applied research into millipede control. Even of the Portugese millipede Ommatoiulus moreletii basic topics have not been addressed, such as the (Diplopoda: Iulidae) in Australia. Australian relative importance of millipedes in nutrient Journal of Ecology 10: 249-259. turnover in forest ecosystems. Burt, J.. 1 984. Report on the research ofthe pincushion millipede, Unixenus mjobergi (Verhoeff, 1924), at Conclusions Tom Price, in the Pilbara of Western Australia. Western Australian Department of Agriculture. The taxonomic framework of millipede orders Golovatch, S.I., 1986. The first Chordeumatida and families in Australia is fairly well known, (Diplopoda) from Tasmania, with the description but there is still much work needed below family of a new genus and three new species. Zoolog- level. The distributions of most species-level isches Jahrbuch, Svstematisches Teil 113: 251- taxa are very poorly known, with the exception 264. of those in the Tasmanian fauna. Basic infor- Griffin, T.T. and Bull, CM.. 1995. Interactions between introduced mation of this kind is required in order to make and native millipede species in South informed decisions about the conservation Australia. Australian Journal ofZoology 43: 129-140. status of Australian millipedes, and at this point Harvey, M.S. and Yen, A.L., 1989. Worms to Wasps: in time such assessment is quite impossible. An Illustrated Guide to Australia's Terrestrial Nevertheless, millipedes have the potential to be Invertebrates. Oxford University Press: quite useful indicator taxa. In many forest and Melbourne. woodland habitats they are common and con- Hoffman, R.. 1979. Classification of the Diplopoda. spicuous, diverse, easily collected and sensitive Museum d'Histoire Naturelle: Geneve. to environmental perturbations. Given the taxo- Hoffman, R., 1982. Diplopoda. Pp. 689-724 in: nomic impediments involved, however, and the Parker, S.P. (ed.). Synopsis and classification of living organisms. McGraw-Hill: almost complete lack of knowledge of their New York. Hoffman, R.L., 1994. Studies on spiroboloid mil- general biology, millipedes cannot at this stage lipeds. XVIII. Speleostrophus nesiotes, the first be recommended as 'environmental indicators' known spiroboloid milliped, from Barrow Island, in Australia. Western Australia (Diplopoda: Pachybolidae: Trigoniulinae). Myriapodologica 3(3): 1*9-24. Acknowledgments Humphreys, W.F. and W.A. Shear, 1993. Troglobitic millipedes (Diplopoda: Paradoxomatidae) from I am grateful to the collection managers and semi-arid Cape Range, Western Australia: sys- curators of myriapod collections at museums tematics and biology. Invertebrate 7: around Australia for making material available 73-195. DIVERSITY AND BIOGEOGRAPHY OF AUSTRALIAN MILLIPEDES 561

Jeekcl, C.A.W., 1981. Australian Expedition 1980; genus Peterjohnsia Mauries, with the description legit C.A.W. Jeekel and A.M. Jeekel-Rijvers. List of a new species from Tasmania (Diplopoda, of collecting stations, together with general notes Chordeumatida, Peterjohnsiidae). Invertebrate on the distributions of millipedes in eastern Aus- Taxonomy 8: 535-544. tralia and Tasmania. Verslagen en Technisehe Shear, W.A. and Mesibov, R., 1995. Australian chor- Gegevens, Instituut voor Taxonomische Zoologie, deumatidan millipeds. II. A new species of Regi- Universitat Amsterdam 30: 1-59. naterreuma Mauries from Tasmania (Diplopoda, Koch, L.E., 1985. Pincushion millipedes (Diplopoda: Chordeumatida, Metopidiotrichidae). Myriapo- Polyxenida): Their aggregations and identity in dologka 3(8): 71-77. Western Australia. Western Australian Naturalist Shulte, F., 1989. The association between Rhabditis 16(2/3): 30-31. neeromena Sudhaus & Shulte, 1989 (Nematoda: Mauries, J-P., 1987. Craspedosomid millipedes dis- Rhabditidae) and native and introduced milli- 82- covered in Australia: Reginaterreuma, Neocam- pedes in South Australia. Nematologiea 35: brisoma and Peterjohnsia, new genera (Myria- 89. poda: Diplopoda: Craspedosomida). Memoirs of Verhoeff, K.W., 1924. Results of Dr. E. Mjoberg's the Queensland Museum 25: 107-133. scientific expeditions to Australia 1910-1913, 34. Shear, W.A. and Mesibov, R.,1994. Australian chor- : Diplopoda. Arkivfoer Zoologi 16(5): deumatidan millipeds. I. New observations on the 1-142.