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Rowe IS05034.Qxd CSIRO PUBLISHING www.publish.csiro.au/journals/is Invertebrate Systematics, 2006, 20, 527–556 Morphological and systematic study of the tribe Australiosomatini (Diplopoda:Polydesmida:Paradoxosomatidea:Paradoxosomatidae) and a revision of the genus Australiosoma Brölemann Melissah RoweA,C and Petra SierwaldB ADepartment of Ecology and Evolution, University of Chicago, 1101 East 57th Street, Chicago, Illinois 60637, USA. BZoology, Field Museum of Natural History, 1400 S Lake Shore Drive, Chicago, IL 60605, USA. CCorresponding author. Email: [email protected] Abstract. The collection of several paradoxosomatid species in the context of ecological studies prompted an investigation into the morphology and species-level characteristics of Australian millipedes in the tribe Australiosomatini Brölemann, 1916 (Polydesmida:Paradoxosomatidae). Three new species are described: Akamptogonus caragoon, sp. nov., Australiosoma fulbrighti, sp. nov. and Australiosoma combei, sp. nov. Notes or re-descriptions are provided for nine additional species belonging to the tribe. Scanning electron microscopy was utilised to examine details of the antennal sensory fields, the fifth sternite lamella and associated pores. The presence of the fifth sternite lamella in adult males is considered a synapomorphy for the family Paradoxosomatidae, whereas the prominent tubercle on the first femur in males (adenostyle) represents a synapomorphy of the subfamily Australiosomatinae. With the description of two new species in the genus Australiosoma Brölemann, 1913 a revision of the genus was undertaken with the purpose of constructing a species- level phylogeny. The most commonly described and utilised species-specific characteristics were scored in a data matrix and analysed using PAUP. The analysis resulted in a single, fully resolved tree of the following structure: Hoplatria clavigera ((A. clavigerum, A. inusitatum) (((A. rainbowi, A. nodulosum) A. michelseni) (A. laminatum (A. combei, A. fulbrighti))). Additional keywords: Australia, millipede. Introduction Systematic history of the subfamily Australiosomatinae The millipede family Paradoxosomatidae Daday, 1889, with Brölemann (1913) first described the genus Australiosoma ~142 genera and more than 560 described species, the (placed in the Polydesmoidea Pocock, 1894) based on the type largest family of the Diplopoda (Hoffman 1980, 1982), has species A. rainbowi Brölemann, 1913, and included a total of a recorded natural range covering all continents excluding five species in this taxon. The group was split into two sub- North America and Antarctica. Currently the family is genera based upon the number of gonopod branches (two organised into three subfamilies (Jeekel 1968: fig. 1; branches in the subgenus Dicladosoma Brölemann, 1913 and Hoffman 1980, 1999), two of which are known from three branches in Australiosoma sensu stricto). Brölemann Australia, the Paradoxosomatinae Daday, 1889 and (1916) argued that the endemic Australian Paradoxosomatidae Australiosomatinae Brölemann, 1916. All known endemic are distinct from those of other countries, implying that they Australian Paradoxosomatidae are contained within the sub- form a monophyletic group, and arranged these species into family Australiosomatinae. Although some species from the two tribes, the Australiosomatini (as Australiosomini) and subfamily Paradoxosomatinae have successfully invaded Antichiropodini (as Antichiropini). At that time, the glasshouses worldwide, e.g. Oxidus gracilis C. L. Koch, Australiosomatini (Brölemann, 1916: 540) contained the 1847, no species from this subfamily are known to be genera Australiosoma and Helicorhabdosoma Brölemann, endemic to Australia. Members of two tribes in the sub- 1916 (now Helicorthomorpha Attems, 1914) and the family Australiosomatinae, the Australiosomatini and Antichiropodini contained the genera Antichiropus Attems, Antichiropodini Brölemann, 1916, are distributed through- 1911 and Haplogonosoma Brölemann, 1916 (Brölemann out Australia, including Tasmania, with some 94 nominal 1916: 539). Attems (1914: 186, couplet 9 in the key; 1929: species assigned to 36 genera (Australiosomatini 20 genera; 262) recognised that the Australian paradoxosomatid genera Antichiropodini 16 genera). can be separated from other paradoxosomatids by the adeno- © CSIRO 2006 10.1071/IS05034 1445-5226/06/050527 528 Invertebrate Systematics M. Rowe and P. Sierwald style (a tubercle, see Fig. 13) on the femur of the first leg in solenomere of medium length. The Antichiropodini were males. Verhoeff (1924: 13, 1928a: 1601, 1941: 5) noted that distinguished by a reduced or absent tibiotarsus, a long the femoral adenostyle on the first leg pair of males occurs in femorite and a very long solenomere. Jeekel (1968) subdi- all then known genera of the Australiosomini and vided the Australiosomatini into five groups based on Antichiropodini. He characterised the Australiosomatini (as gonopod characters, such as the number of branches and Australiosominae) by the free strong solenomere and the their relative lengths and positions (see Jeekel 1968: fig. 1). strongly developed tibiotarsus. Jeekel (1964: 108) noted that Jeekel hypothesised about the primitive condition of the the adenostyle is also present in Dendrogonopus Jeekel, 1964 gonopods in Australiosomatini, including the reductive ten- and thus also occurs in the third tribe of the subfamily dencies of the femorite, the enlargement of the femoral Australiosomatinae, tribe Aschistodesmini (Fig. 1). process and bifurcation of the tibiotarsus. Jeekel (1968), following Brölemann and Verhoeff, sepa- Currently, the Australiosomatini are characterised by the rated the tribes employing the following characters: following combination of characters: a short or somewhat Australiosomatini: tibiotarsus present, femorite short, elongate gonopod prefemur, thick and rather short to almost Australiosomatini Australia Australiosomatinae Australia P Antichiropodini a Aschistodesmini New Guinea r a Alogolykinae Alogolykini Myanmar, China d o Polydrepanini India x o s Eviulisomatini Africa o Eroonsomatini Italy, Yugoslavia m Graphisternini Peru a Eustrongylosomatini New Guinea, Philippines t Cnemodesmini Africa i Sulciferini Indochina d Orthomorphini (incl. ‘Hylomini’) SE Asia a Paradoxosomatinae Strongylosomatini Golovatch 1984 India e Chamberlinini Taiwan Tectoporini SE Asia Tonkinosomatini Indochina Nedyopodini SE Asia Sundanini mainly Indonesia Xanthodesmini Pakistan, Africa Paradoxosomatini Europe, middle East Catharosomatini South America Fig. 1. Proposed relationships within the family Paradoxosomatidae, from Jeekel (1968) and Hoffman (1980), modified to incorporate classification changes by Golovatch (1984) and Golovatch and Enghoff (1994). Australian Paradoxosomatidae Invertebrate Systematics 529 entirely absent femorite and a strongly developed Materials and methods solenomere (Jeekel 1968, 1987). In many genera, the Specimens examined included millipedes collected during ecological gonopod femorite is provided with a strong femoral process studies and type material borrowed from museums. Material was not arising from the anterior side of the femorite. In addition to obtained by sorting though the extensive backlog at various Australian these gonopod features, the paranota are moderately devel- institutions (see Sierwald and Reft 2004); such a task was outside the scope of this study. Millipedes were hand-collected in the Sydney Basin oped. The various distinguishing characters employed by region during May 2000. Collection times were synchronised with previous authors demonstrate that the Australiosomatini are observed peak activity periods of 06:00 hours to 12:00 hours, with most not defined by unequivocal apomorphies. success on overcast days or during light rain. Specimens were located Representatives of the tribe have been collected from all in leaf litter, fallen logs and on trees during random walk searches. regions within Australia, with most attention to the Specimens were euthanised by cooling and then preserved in 70% Queensland (Jeekel 1987) and South Australian (Jeekel ethanol in glass vials of appropriate size, cotton plugged and placed in parfait jars filled with 70% ethanol. 1985) fauna. The tribe currently includes 20 genera and Specimens were examined using light and scanning electron 56 species (the number of species for each genus are listed microscopy (SEM). Scanning electron micrographs of the gonopods below in italics): Akamptogonus Attems, 1914, 1; Archi- did not reveal features, such as setae or pores, that were not already cladosoma Jeekel, 1984, 1; Australiosoma Brölemann, 1913, observed using light microscopy. However, the SEM image imparts a 6; Boreohesperus Shear, 1992, 1; Cladethosoma three-dimensional view of the structures, whereas the line drawings indicate overlaying structures. Additional characteristics explored in Chamberlin, 1920, 7; Dicladosoma Brölemann, 1913, 1; the current study include the cuticle structure, spiracles, gnathochilar- Dicladosomella Jeekel, 1982, 1; Gigantowales Verhoeff, ium, antennae, epiproct and in males, the lamella on the fifth sternite, 1937, 2; Heterocladosoma Jeekel, 1968, 7; Hoplatessara the gonopore and legs. Gonopods were dissected and drawn with the aid Verhoeff, 1928, 8; Hoplatria Verhoeff, 1941, 1; Isoclado- of a Leica MZ12 microscope and a camera lucida. Pencil drawings were soma Jeekel, 1984, 3; Myallosoma Verhoeff, 1928, 2; Onco- digitised using Adobe Illustrator. Specimen samples for electron microscopy were
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