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Araneae: Mygalomorphae: Actinopodidae: Missulena) from the Pilbara Region, Western Australia

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Araneae: Mygalomorphae: Actinopodidae: Missulena) from the Pilbara Region, Western Australia Zootaxa 3637 (5): 521–540 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3637.5.2 http://zoobank.org/urn:lsid:zoobank.org:pub:447D8DF5-F922-4B3A-AC43-A85225E56C57 New species of Mouse Spiders (Araneae: Mygalomorphae: Actinopodidae: Missulena) from the Pilbara region, Western Australia DANILO HARMS1, 2, 3, 4 & VOLKER W. FRAMENAU1, 2, 3 1 School of Animal Biology, The University of Western Australia, 35 Stirling Highway, Crawley, Western Australia 6009, Australia. 2 Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia. 3 Phoenix Environmental Sciences Pty Ltd, 1/511 Wanneroo Road, Balcatta, Western Australia 6021, Australia. 4 Corresponding author. E-mail: [email protected] Abstract Two new species of Mouse Spiders, genus Missulena, from the Pilbara region in Western Australia are described based on morphological features of males. Missulena faulderi sp. nov. and Missulena langlandsi sp. nov. are currently known from a small area in the southern Pilbara only. Mitochondrial cytochrome c oxidase subunit I (COI) sequence divergence failed in clearly delimiting species in Missulena, but provided a useful, independent line of evidence for taxonomic work in addition to morphology. Key words: taxonomy, systematics, barcoding, mitochondrial DNA, short-range endemism, Actinopus, Plesiolena Introduction The Actinopodidae Simon, 1892 is a small family of mygalomorph spiders with a Gondwanan distribution that includes three genera: Actinopus Perty, 1833 (27 species), Missulena Walckenaer, 1805 (11 species) and Plesiolena Goloboff & Platnick, 1987 (two species). Actinopus and Plesiolena are known only from South and Central America (Platnick 2012). In contrast, Missulena Walckenaer, 1805 includes 10 species from Australia and one species, M. tussulena Goloboff, 1994, from Chile. Australian species of Missulena, commonly known as Mouse Spiders, are medium-sized spiders with a steeply elevated cephalic region and a wide eye group (Fig. 1A). Some species have received public attention due to the toxicity of their venom that has been found to be biochemically similar to that of Australian Funnel-web Spiders (family Hexathelidae Simon, 1892) (Isbister 2004; Herzig et al. 2008; Rash et al. 2000). Missulena insignis (O.P.- Cambridge, 1877), M. occatoria Walckenaer, 1805 and M. reflexa Rainbow & Pulleine, 1918 are known for conspicuous red fangs and cephalic areas in males, although many described species, e.g. M. bradleyi Rainbow, 1914, M. dipsaca Faulder, 1995, M. granulosa (O. Pickard-Cambridge, 1869), M. rutraspina Faulder, 1995 and M. torbayensis Main, 1996 are uniformly dark brown or black in colour (Main 1956, 1996). The taxonomy of Australian Missulena was first reviewed by Womersley (1943), who recognised and diagnosed six species; only four of these are known from both male and female specimens. Main (1985) summarised the current knowledge of Missulena and proposed several taxonomic changes. Shortly after, Faulder (1995) published descriptions of two widespread new species of Missulena: M. dipsaca and M. rutraspina. More recently, Main (1996) named M. torbayensis from southwestern Western Australia and noted that “it is apparent from my field observations and an abundance of specimens (mainly males) in museums that there are many undescribed species.” Recent large-scale biological surveys in the Carnarvon Basin (Main et al. 2000) and Pilbara region (Durrant et al. 2010) of Western Australia have added substantial material of Missulena, in particular males, to the collection of the Western Australian Museum (Fig. 2). Our work is based on this material and has two aims: first, we describe males of two morphologically distinct species of Missulena from the Pilbara region. Second, we conduct a Accepted by R. Raven: 11 Feb. 2013; published: 15 Apr. 2013 521 molecular analyses using a fragment of the cytochrome oxidase c subunit I (COI) gene from a selection of Missulena specimens from Western Australia to provide an additional and independent line of evidence in the taxonomic process, and to review the suitability of this marker for single-gene barcoding of Missulena species. Material and methods Morphology. Specimens used for morphological examination were preserved in 75% ethanol and examined using a Leica M80 stereomicroscope. Digital images were taken using a Leica DFC295 digital camera attached to a Leica M205C stereomicroscope controlled by the Leica Application Suite Version 2.5. Distribution maps were produced using the Open Source software package Quantum GIS Version 1.7.4 Wroclaw (www.qgis.org; accessed 30 May 2012). Climate data for the Pilbara bioregion were retrieved from the Australian Government, Bureau of Meteorology (online under www.bom.gov.au/climate/averages/tables) and refer to the airport of Newman. Measurements are expressed in millimetres and were taken in dorsal view, except those of labium, sternum, coxae, trochanters and spigots, which were taken in ventral view. Total lengths were taken in dorsal view and exclude the chelicerae and spinnerets. The number of teeth on the claws is given as the formula “leg number: number of teeth of lateral claws / number of teeth of median claw”. The leg formula is given as the order of the leg lengths from longest to shortest. The leg “index” is given here as the leg length divided by carapace length and indicates the ratio of leg lengths versus carapace. Measurements of the eye region refer to the distance between the two most separate eyes in a row; those of the ocular quadrangle (OQ) to the quadrangle limited by the four lateral eyes, and those of the median ocular quadrangle (MOQ) to the distance between the four median eyes. The term “rasps” refers to the presence of short but strong conical spines at the patellae of legs I–IV. The occurrence of such spines on patella I has been suggested as a potential synapomorphy for Missulena species (Goloboff & Platnick 1987). Our species hypotheses are based only on males, as are those established by previous authors (Womersley 1943, Faulder 1995, Main 1996). All specimens are lodged at the WA Museum (WAM). Molecular methods. We sequenced 22 specimens of Missulena from Western Australia (Figs 2A, B, 3; Table 1). Reference specimens were from five additional bioregions of Western Australia (Thackway & Cresswell 1995): Avon Wheatbelt, Jarrah Forest, Gascoyne, Murchison and Ord Victoria Plain (Figs 2A, 3). Each specimen sequenced as part of this study is denoted by a superscript DNA code in the Material Examined lists, following Harvey et al. (2012). We also sequenced two putatively undescribed species of Conothele Thorell, 1878 (Ctenizidae Thorell, 1887) and retrieved an additional sequence of Euagrus chisoseus Gertsch, 1939 (Dipluridae Simon, 1889) from Genbank for use as an ultimate outgroup (Table 1). All specimens were sequenced for variation at the mitochondrial cytochrome c oxidase subunit I (COI) gene which is preferred here over other potential barcoding genes because of the general lack of introns and infrequency of nucleotide polymorphisms, high inter- and intra-specific variability, and unproblematic amplification using standard methodology (Folmer et al. 1994, Hebert et al. 2003). Genomic DNA was extracted from leg muscle tissue using the QIAGEN DNeasy Blood and Tissue Kit for animal tissues (QIAGEN Inc., Valencia, CA). Polymerase chain reaction (PCR) amplification of an 827 bp fragment of the COI gene was achieved using the primers LCO1490 (Folmer et al. 1994) and HCOoutout (Prendini et al. 2005). PCR products were sequenced for both DNA strands at the AGRF node in Perth (Western Australian Institute of Medical Research, WA, Australia). Sequences were checked by eye, edited and aligned using the program Geneious Version 5.6 (Drummond et al. 2011). Tree building. Two Bayesian phylogenetic analyses were executed using MrBayes Version 3.2.1 for Macintosh (Huelsenbeck & Ronquist 2001; Ronquist & Huelsenbeck 2003) applying 1) three partitions according to codon position, and 2) one partition across all data. MrModeltest Version 3.7 (Posada & Crandall 1998) was used to estimate the appropriate model of nucleotide substitution under an Akaike Information Criterion (AIC) framework. A general time reversible model (GTR) (Rodríguez et al. 1990) with a class of invariable sites (I) and gamma distributed rate heterogeneity (G) (Yang 1996) was invoked for the unpartitioned data and codon positions 1 and 2. A GTR model with gamma distributed rate heterogeneity (GTR+G) was suggested for the third codon position. Four Markov chain Monte Carlo (MCMC) chains were run for 40,000,000 generations to achieve convergence of the independent runs, sampling every 1,000 generations and discarding the first 25% of sampled 522 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU trees as ‘burnin’. FigTree Version 1.3.1 (Rambaut 2009) was used to visualise and edit tree files, and Tracer Version 1.5 (Rambaut & Drummond 2009) was used to check parameter values and convergence of the MCMC chains. To infer the degree of genetic similarity between the sequenced specimens, we finally calculated a distance matrix in Geneious for the sequenced Missulena specimens using the edited and aligned DNA sequence data. Abbreviations AME = anterior median eyes, ALE = anterior lateral eyes, MOQ = median ocular quadrangle, OC = ocular quadrangle, PME = posterior
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