Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

A revision of the textricellin spider genus Raveniella (Araneae : Araneoidea : Micropholcommatidae): exploring patterns of phylogeny and biogeography in an Australian biodiversity hotspot

Michael G. Rix A B C , Mark S. Harvey B A and J. Dale Roberts A

A School of Animal Biology M092, The University of Western Australia, 35 Stirling Highway, Crawley, Perth, WA 6009, Australia.

B Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welsphool D.C., Perth, WA 6986, Australia.

C Corresponding author. Email:

Invertebrate Systematics 24(3) 209-237
Submitted: 4 December 2009  Accepted: 12 June 2010   Published: 30 August 2010


South-western Western Australia is a biodiversity hotspot, with high levels of local endemism and a rich but largely undescribed terrestrial invertebrate fauna. Very few phylogeographic studies have been undertaken on south-western Australian invertebrate taxa, and almost nothing is known about historical biogeographic or cladogenic processes, particularly on the relatively young, speciose Quaternary sand dune habitats of the Swan Coastal Plain. Phylogeographic and taxonomic patterns were studied in textricellin micropholcommatid spiders belonging to the genus Raveniella Rix & Harvey. The Micropholcommatidae is a family of small spiders with a widespread distribution in southern Western Australia, and most species are spatially restricted to refugial microhabitats. In total, 340 specimens of Raveniella were collected from 36 surveyed localities on the Swan Coastal Plain and 17 non-Swan Coastal Plain reference localities in south-western Western Australia. Fragments from three nuclear rRNA genes (5.8S, 18S and ITS2), and one mitochondrial protein-coding gene (COI) were used to infer the phylogeny of the genus Raveniella, and to examine phylogeographic patterns on the Swan Coastal Plain. Five new species of Raveniella are described from Western Australia (R. arenacea, sp. nov., R. cirrata, sp. nov., R. janineae, sp. nov., R. mucronata, sp. nov. and R. subcirrata, sp. nov.), along with a single new species from south-eastern Australia (R. apopsis, sp. nov.). Four species of Raveniella were found on the Swan Coastal Plain: two with broader distributions in the High Rainfall and Transitional Rainfall Zones (R. peckorum Rix & Harvey, R. cirrata); and two endemic to the Swan Coastal Plain, found only on the western-most Quindalup dunes (R. arenacea, R. subcirrata). Two coastally restricted species (R. subcirrata, R. janineae) were found to be morphologically cryptic but genetically highly distinct, with female specimens morphologically indistinguishable from their respective sister-taxa (R. cirrata and R. peckorum). The greater Perth region is an important biogeographic overlap zone for all four Swan Coastal Plain species, where the ranges of two endemic coastal species join the northern and south-western limits of the ranges of R. peckorum and R. cirrata, respectively. Most species of Raveniella were found to occupy long, highly autapomorphic molecular branches exhibiting little intraspecific variation, and an analysis of ITS2 rRNA secondary structures among different species of Raveniella revealed the presence of an extraordinary hypervariable helix, ranging from 31 to over 400 nucleotides in length.


Bond J. E. Stockman A. K. 2008 An integrative method for delimiting cohesion species: finding the population–species interface in a group of Californian trapdoor spiders with extreme genetic divergence and geographic structuring. Systematic Biology 57 628 646 doi:10.1080/10635150802302443

Bond J. E. Hedin M. C. Ramírez M. G. Opells B. D. 2001 Deep molecular divergence in the absence of morphological and ecological change in the Californian coastal dune endemic trapdoor spider Aptostichus simus. Molecular Ecology 10 899 910 doi:10.1046/j.1365-294X.2001.01233.x

Brown R. W. (1956). ‘Composition of Scientific Words: A Manual of Methods and a Lexicon of Materials for the Practice of Logotechnics.’ (Smithsonian Books: Washington, DC.)

Browne-Cooper R. Maryan B. 1992 Notes on the status of the skink lizard Ctenotus lancelini on Lancelin Island. Western Australian Naturalist 19 63 65

Burbidge A. A. Kutchling G. 2007 The western swamp tortoise – 50 years on. Landscope 22 24 29

Coleman A. W. 2007 Pan-eukaryote ITS2 homologies revealed by RNA secondary structure. Nucleic Acids Research 35 3322 3329 doi:10.1093/nar/gkm233

Crews S. C. Hedin M. 2006 Studies of morphological and molecular phylogenetic divergence in spiders (Araneae: Homalonychus) from the American southwest, including divergence along the Baja California Peninsula. Molecular Phylogenetics and Evolution 38 470 487 doi:10.1016/j.ympev.2005.11.010

Dadour I. R. Johnson M. S. 1983 Genetic differentiation, hybridization and reproductive isolation in Mygalopsis marki Bailey (Orthoptera: Tettigoniidae). Australian Journal of Zoology 31 353 360 doi:10.1071/ZO9830353

Department of the Environment Water, Heritage and the Arts (2009). Ctenotus lancelini. In ‘Species Profile and Threats Database, Department of the Environment, Water, Heritage and the Arts, Canberra’. Available at [Accessed 26 January 2009].

Edward K. L. Harvey M. S. 2010 A review of the Australian millipede genus Atelomastix (Diplopoda: Spirostreptida: Iulomorphidae). Zootaxa 2371 1 63

Edwards D. L. Roberts J. D. Keogh J. S. 2007 Impact of Plio-Pleistocene arid cycling on the population history of a southwestern Australian frog. Molecular Ecology 16 2782 2796 doi:10.1111/j.1365-294X.2007.03374.x

Edwards D. L. Roberts J. D. Keogh J. S. 2008 Climatic fluctuations shape the phylogeography of a mesic direct-developing frog from the south-western Australian biodiversity hotspot. Journal of Biogeography 35 1803 1815 doi:10.1111/j.1365-2699.2008.01927.x

Felsenstein J. 1985 Confidence limits on phylogenies – an approach using the bootstrap. Evolution 39 783 791 doi:10.2307/2408678

Forster R. R. 1959 The spiders of the family Symphytognathidae. Transactions of the Royal Society of New Zealand 86 269 329

Forster R. R. Platnick N. I. 1981 A textricellid spider from Chile (Araneae, Textricellidae). Bulletin of the American Museum of Natural History 170 263 270

Forster R. R. Platnick N. I. 1984 A review of the archaeid spiders and their relatives, with notes on the limits of the superfamily Palpimanoidea (Arachnida, Araneae). Bulletin of the American Museum of Natural History 178 1 106

Harvey M. S. 1996 The biogeography of Gondwanan pseudoscorpions (Arachnida). Revue Suisse de Zoologie vol. hors série 255 264

Harvey M. S. 2002 a Short-range endemism among the Australian fauna: some examples from non-marine environments. Invertebrate Systematics 16 555 570 doi:10.1071/IS02009

Harvey M. S. 2002 b A new species of Austrarchaea (Araneae: Archaeidae) from Western Australia. Records of the Western Australian Museum 21 35 37

Harvey M. S. Waldock J. M. Guthrie N. A. Durrant B. J. McKenzie N. L. 2004 Patterns of composition of ground-dwelling araneomorph spider communities in the Western Australian Wheatbelt. Records of the Western Australian Museum Supplement No. 67 257 291

Harvey M. S. Berry O. Edward K. L. Humphreys G. 2008 Molecular and morphological systematics of hypogean schizomids (Schizomida: Hubbardiidae) in semiarid Australia. Invertebrate Systematics 22 167 194 doi:10.1071/IS07026

Hedin M. C. 1997 Speciational history in a diverse clade of habitat-specialized spiders (Araneae: Nesticidae: Nesticus): inferences from geographic-based sampling. Evolution 51 1929 1945 doi:10.2307/2411014

Hendrixson B. E. Bond J. E. 2005 Testing species boundaries in the Antrodiaetus unicolor complex (Araneae: Mygalomorphae: Antrodiaetidae): “paraphyly” and cryptic diversity. Molecular Phylogenetics and Evolution 36 405 416 doi:10.1016/j.ympev.2005.01.021

Hickman V. V. 1944 On some new Australian Apneumonomorphae with notes on their respiratory system. Papers and Proceedings of the Royal Society of Tasmania 1944 179 195

Hickman V. V. 1945 A new group of apneumone spiders. Transactions of the Connecticut Academy of Arts and Sciences 36 135 148

Hill A. Johnson M. S. Merrifield H. 1983 An electrophoretic and morphological examination of Bothriembryon kendricki (Pulmonata: Bulimulidae), a new species previously considered conspecific with B. bulla (Menke). Australian Journal of Zoology 31 227 242 doi:10.1071/ZO9830227

Hong-Chun P. Kai-Ya Z. Da-Xiang S. Yang Q. 2004 Phylogenetic placement of the spider genus Nephila (Araneae: Araneoidea) inferred from rRNA and MaSp1 gene sequences. Zoological Science 21 343 351 doi:10.2108/zsj.21.343

Hopper S. D. Gioia P. 2004 The Southwest Australian Floristic Region: evolution and conservation of a global hot spot of biodiversity. Annual Review of Ecology Evolution and Systematics 35 623 650 doi:10.1146/annurev.ecolsys.35.112202.130201

Hopper S. D. , Harvey M. S. , Chappill J. A. , Main A. R. , and Main B. Y. (1996). The Western Australian biota as Gondwanan heritage – a review. In ‘Gondwanan Heritage. Past Present and Future of the Western Australian Biota’. (Eds S. D. Hopper, J. A. Chappill, M. S. Harvey and A. S. George.) pp. 1–46. (Surrey Beatty & Sons: Chipping Norton, NSW.)

Huelsenbeck J. P. Ronquist F. 2001 MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics (Oxford, England) 17 754 755 doi:10.1093/bioinformatics/17.8.754

Iredale T. 1939 A review of the land Mollusca of Western Australia. Records of the Western Australian Museum and Art Gallery 2 1 88

Keller A. Schleicher T. Schultz J. Müller T. Dandekar T. Wolf M. 2009 5.8S–28S rRNA interaction and HMM-based ITS2 annotation. Gene 430 50 57 doi:10.1016/j.gene.2008.10.012

Kendrick G. W. Wyrwoll K.-H. Szabo B. J. 1991 Pliocene–Pleistocene coastal events and history along the western margin of Australia. Quaternary Science Reviews 10 419 439 doi:10.1016/0277-3791(91)90005-F

Littlejohn M. J. 1957 A new species of frog of the genus Crinia. Western Australian Naturalist 6 18 23

Longhorn S. J. Nicholas M. Chuter J. Vogler A. P. 2007 The utility of molecular markers from non-lethal DNA samples of the CITES II protected “tarantula” Brachypelma vagans (Araneae, Theraphosidae). The Journal of Arachnology 35 278 292 doi:10.1636/S05-62.1

Main B. Y. 1996 Terrestrial invertebrates in south-west Australian forests: the role of relict species and habitats in reserve design. Journal of the Royal Society of Western Australia 79 277 280

Main B. Y. (1999). Biological anachronisms among trapdoor spiders reflect Australia’s environmental changes since the Mesozoic. In ‘The Other 99%. The Conservation and Biodiversity of Invertebrates’. (Eds W. Ponder and D. Lunney.) pp. 236–245. (Transactions of the Royal Zoological Society of New South Wales: Mosman, NSW.)

Main B. Y. Harvey M. S. Waldock J. M. 2002 The distribution of the Western Australian pill millipede, Cynotelopus notabilis Jeekel (Sphaerotheriiidae). Records of the Western Australian Museum 20 383 385

Mast A. R. Jones E. H. Havery S. P. 2005 An assessment of old and new DNA sequence evidence for the paraphyly of Banksia with respect to Dryandra (Proteaceae). Australian Systematic Botany 18 75 88 doi:10.1071/SB04015

Mathews D. H. Disney M. D. Childs J. L. Schroeder S. J. Zuker M. Turner D. H. 2004 Incorporating chemical modification constraints into a dynamic programming algorithm for prediction of RNA secondary structure. Proceedings of the National Academy of Sciences of the United States of America 101 7287 7292 doi:10.1073/pnas.0401799101

Moir M. L. Brennan K. E. C. Harvey M. S. 2009 Diversity, endemism and species turnover of millipedes within the south-western Australian global biodiversity hotspot. Journal of Biogeography 36 1958 1971

Morgan M. J. Roberts J. D. Keogh J. S. 2007 Molecular phylogenetic dating supports an ancient endemic speciation model in Australia’s biodiversity hotspot. Molecular Phylogenetics and Evolution 44 371 385 doi:10.1016/j.ympev.2006.12.009

Murienne J. Pellens R. Budinoff R. B. Wheeler W. C. Grandcolas P. 2008 Phylogenetic analysis of the endemic New Caledonian cockroach Lauraesilpha. Testing competing hypotheses of diversification. Cladistics 24 802 812 doi:10.1111/j.1096-0031.2008.00204.x

Myers N. Mittermeier R. A. Mittermeier C. G. da Fonseca G. A. B. Kent J. 2000 Biodiversity hotspots for conservation priorities. Nature 403 853 858 doi:10.1038/35002501

Nylander J. A. A. (2004). ‘MrModeltest Version 2.’ Available at [Accessed on 26 January 2009].

Posada D. Crandall K. A. 1998 Modeltest: testing the model of DNA substitution. Bioinformatics (Oxford, England) 14 817 818 doi:10.1093/bioinformatics/14.9.817

Read K. Keogh J. S. Scott I. A. W. Roberts J. D. Doughty P. 2001 Molecular phylogeny of the Australian frog genera Crinia, Geocrinia, and allied taxa (Anura: Myobatrachidae). Molecular Phylogenetics and Evolution 21 294 308 doi:10.1006/mpev.2001.1014

Reid A. 2002 Western Australian Onychophora (Peripatopsidae): a new genus, Kumbadjena, for a southern species-complex. Records of the Western Australian Museum 21 129 155

Rix M. G. 2006 Systematics of the Australasian spider family Pararchaeidae (Arachnida: Araneae). Invertebrate Systematics 20 203 254 doi:10.1071/IS05032

Rix M. G. 2008 A new species of Micropholcomma (Araneae: Araneoidea: Micropholcommatidae) from Western Australia. Records of the Western Australian Museum 24 343 348

Rix M. G. Harvey M. S. 2010 The spider family Micropholcommatidae (Arachnida: Araneae: Araneoidea): a relimitation and revision at the generic level. ZooKeys 36 1 321 doi:10.3897/zookeys.36.306

Rix M. G. Harvey M. S. Roberts J. D. 2008 Molecular phylogenetics of the spider family Micropholcommatidae (Arachnida: Araneae) using nuclear rRNA genes (18S and 28S). Molecular Phylogenetics and Evolution 46 1031 1048 doi:10.1016/j.ympev.2007.11.001

Ronquist F. Huelsenbeck J. P. 2003 MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics (Oxford, England) 19 1572 1574 doi:10.1093/bioinformatics/btg180

Rozen S. , and Skaletsky H. J. (2000). Primer3 on the WWW for general users and for biologist programmers. In ‘Bioinformatics Methods and Protocols: Methods in Molecular Biology’. (Eds S. Krawetz and S. Misener.) pp. 365–386. (Humana Press: Totowa, NJ.)

Sanmartín I. Ronquist F. 2004 Southern hemisphere biogeography inferred by event-based models: plant versus animal patterns. Systematic Biology 53 216 243 doi:10.1080/10635150490423430

Schnare M. N. Damberger S. H. Gray M. W. Gutell R. R. 1996 Comprehensive comparison of structural characteristics in eukaryotic cytoplasmic large subunit (23 S-like) ribosomal RNA. Journal of Molecular Biology 256 701 719 doi:10.1006/jmbi.1996.0119

Schultz J. Wolf M. 2009 ITS2 sequence-structure analysis in phylogenetics: a how-to manual for molecular systematics. Molecular Phylogenetics and Evolution 52 520 523 doi:10.1016/j.ympev.2009.01.008

Seddon G. (1972). ‘Sense of Place: A Response to an Environment, the Swan Coastal Plain, Western Australia.’ (University of Western Australia Press: Perth.)

Semeniuk V. Semeniuk C. A. 2006 Sedimentary fill of basin wetlands, central Swan Coastal Plain, southwestern Australia. Part 2: distribution of sediment types and their stratigraphy. Royal Society of Western Australia 89 185 220

Simon C. Frati F. Beckenbach A. Crespi B. Liu H. Flook P. 1994 Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87 651 701

Sparks J. S. Smith W. L. 2004 Phylogeny and biogeography of the Malagasy and Australasian rainbowfishes (Teleostei: Melanotaenioidei): Gondwanan vicariance and evolution in freshwater. Molecular Phylogenetics and Evolution 33 719 734 doi:10.1016/j.ympev.2004.07.002

Stockman A. K. Bond J. E. 2008 A taxonomic review of the trapdoor spider genus Promyrmekiaphila Schenkel (Araneae, Mygalomorphae, Cyrtaucheniidae, Euctenizinae). Zootaxa 1823 25 41

Swofford D. L. (2002). ‘PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4.0b10.’ (Sinauer Associates: Sunderland, MA.)

Tapsell P. Newsome D. Bastian L. 2003 Origin of yellow sand from Tamala Limestone on the Swan Coastal Plain, Western Australia. Australian Journal of Earth Sciences 50 331 342 doi:10.1046/j.1440-0952.2003.00989.x

Tautz D. Hancock J. M. Webb D. A. Tautz C. Dover G. A. 1988 Complete sequences of the rRNA genes of Drosophila melanogaster. Molecular Biology and Evolution 5 366 376

Thompson J. D. Gibson T. J. Plewniak F. Jeanmougin F. Higgins D. G. 1997 The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25 4876 4882 doi:10.1093/nar/25.24.4876

Wardell-Johnson G. Roberts J. G. 1993 Biogeographic barriers in a subdued landscape: the distribution of the Geocrinia rosea (Anura: Myobatrachidae) complex in south-western Australia. Journal of Biogeography 20 95 108 doi:10.2307/2845743

White T. J. , Bruns T. , Lee S. , and Taylor J. (1990). Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In ‘PCR Protocols: A Guide to Methods and Applications’. (Eds M. Innis, D. Gelfand, J. Swinksy and T. White.) pp. 315–322. (Academic Press: San Diego, CA.)

Wuyts J. De Rijk P. Van de Peer Y. Pison G. Rousseeuw P. De Wachter R. 2000 Comparative analysis of more than 3000 sequences reveals the existence of two pseudoknots in area V4 of eukaryotic small subunit ribosomal RNA. Nucleic Acids Research 28 4698 4708 doi:10.1093/nar/28.23.4698

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