Vink, C. J.; Dupérré, N.; Malumbres-Olarte, J. 2013. Periegopidae (Arachnida: Araneae). Fauna of New Zealand 70: 41 pp.

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C. J. Vink AgResearch, Biocontrol & Biosecurity Private Bag 4749, Christchurch 8140, New Zealand (present address: Canterbury Museum, Rolleston Avenue, Christchurch 8013, New Zealand [email protected]) Entomology Research Museum PO Box 84, Lincoln University, Lincoln 7647, New Zealand

N. Dupérré 341 15ème Rue, Laval, Québec, H7N 1L5, Canada [email protected]

J. Malumbres-Olarte Department of Ecology PO Box 84, Lincoln University, Lincoln 7647, New Zealand [email protected]

Manaaki W h e n u a P R E S S

Lincoln, Canterbury, New Zealand 2013 4 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

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Cataloguing in publication Vink, C. J. (Cornelis Jacob) Periegopidae (Arachnida: Araneae) / C. J. Vink, N. Dupérré and J. Malumbres-Olarte. – Lincoln, Canterbury, N.Z. : Manaaki Whenua Press, 2013. (Fauna of New Zealand, ISSN 0111-5383 (print), ISSN 1179-7193 (online) ; no. 70). ISBN 978-0-478-34740-1 (print) ISBN 978-0-478-34741-8 (online)

I. Dupérré, N. (Nadine) II. Malumbres-Olarte, J. (Jagoba) III. Title. IV. Series. UDC 595.44

Suggested citation: Vink, C. J.; Dupérré, N.; Malumbres-Olarte, J. 2013. Periegopidae (Arachnida: Araneae). Fauna of New Zealand 70: 41 pp.

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Date of publication 7 March 2013

Front cover: Female Periegops suterii Urquhart from Kennedys Bush Reserve, MC (LUNZ 00012718) (Photographer: Bryce McQuillan).

Publication of the Fauna of New Zealand series is supported by Core funding for Crown Research Institutes from the Ministry of Business, Innovation and Employment’s Science and Innovation Group, for the Defining New Zealand’s Land Biota research programme. Fauna of New Zealand 70 ­5­

POPULAR SUMMARY HE WHAKARĀPOPOTOTANGA Class Periegopidae Order Arachnida Family Araneae

Periegopid spiders There are only three species known in the family Perigopi- dae and all are in one genus, Periegops. These rare spiders have only ever been found in relict forests at limited loca- tions in New Zealand (Banks Peninsula, Riccarton Bush, the Aldermen Islands, and East Cape) and Queensland, Australia. Periegopids are only found in forest with a deep leaf litter layer and well-drained soil. They do not build a web, but hunt on the forest floor. Periegopids can be most readily distinguished from other spiders found in New Zealand by having six eyes arranged in three widely spaced diads.

Contributor Cor Vink was born and educated in Christch- Illustration / Whakaahua: Periegops suterii, female urch, New Zealand. He completed a Ph.D. at Lincoln (Illustrator / Kaiwhakaahua: N. Dupérré). University on the taxonomy and systematics of New Zealand Lycosidae, a major part of which was published as a revision in Fauna of New Zealand 44. After complet- Ngā pūngāwere Periegopid ing his thesis he spent nine months at AgResearch as a E toru anake ngā momo e mōhiotia ana o te whānau postdoctoral research fellow investigating the genetics of Perigopidae, ā, nō te huinga kotahi, arā, a Periegops. Kua hymenopteran parasitoids of weevil pests. From 2003 to kitea ēnei pūngāwere onge tonu i ngā toenga ngahere i ngā 2005, Cor was a postdoctoral associate at San Diego State wāhi ruarua nei i Aotearoa (i Horomaka, i te ngahere o Ric- University, U.S.A., where he worked on developing new carton, i ngā moutere Aldermen, me Te Koringa-a-Paoa) molecular markers for inferring deep phylogenetic rela- me Queensland, i Ahitereiria. Kitea ai ēnei Periegopid i te tionships in spiders. At the end of 2005 he returned to New ngahere he hōhonu nei ōna pūranga rau-rākau, i ngā one Zealand and joined the Biosecurity Group at AgResearch, āhua pai te mimititanga wai. Kāore e hanga tukutuku mō Lincoln as a scientist. Cor is especially interested in the rātou, engari ka whakangau i ā rātou kai i te papa tonu o systematics of New Zealand spiders and is the adjunct te ngahere. Ka mōhiotia wawetia te Periegopid, ina wha- curator of spiders at the Entomology Research Museum at karitea ki ētahi katoa o ngā pūngāwere o Aotearoa, nā te Lincoln University. In 2013, Cor was appointed as Curator noho mai o ngā karu e ono, he tawhiti tonu te takoto o ia Natural History at Canterbury Museum, Christchurch. pūruatanga karu i ētahi atu. Translation by Piripi Walker (continued overleaf) Whakatiki 6 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Contributor Nadine Dupérré is from Québec, Canada. She completed a Bachelor of Science at Université de Montréal in 1997. She began illustrating spiders in 1998 and to date has produced over 5000 illustrations. Nadine is probably best known amongst arachnologists for her illustrations in the book “Spiders of North America: an Identification Manual”. Nadine has published on the taxonomy of spiders and has also produced illustrations for publications on beetles and harvestmen. Nadine was a research assistant at the American Museum of Natural History, New York from 2008–2012 where she worked on Oonopidae with Norman Platnick. She is now living in Quito, Ecuador, working on a revision of the genus Agyneta (Linyphiidae) from North America, and is planning to continue her work on Linyphiidae in South America.

Contributor Jagoba Malumbres-Olarte is from the Basque Country, Spain. He completed a Ph.D. at Lincoln University on the ecology and diversity of spider commu- nities in New Zealand native tussock grasslands in 2011. After he completed his thesis, he worked as a teaching

assistant at Lincoln University and Imperial College Lon- Cor Vink don, UK. In the last six years Jagoba has been involved in numerous research projects on diverse topics, including the ecology of spider communities in sand dunes, conser- vation genetics of the Chatham Islands coxella weevil, the systematics of the New Zealand Clubiona, and the invasibility of spiders and plants in New Zealand. Jagoba is particularly interested in ecological and environmental processes that drive speciation and shape the diversity and composition of communities, and is currently developing new projects in this field. Nadine Dupérré Jagoba Malumbres-Olarte Fauna of New Zealand 70 ­7­

ABSTRACT Two species of Periegopidae, both in the genus Periegops Simon, 1893, are found in New Zealand; P. suterii (Urquhart, 1892) and P. keani sp. nov. The genus and both species are described or redescribed, with information on synonymy, type data, material examined, and geographical distribution. Habitus images of adults, illustrations of important morphological features, and distribution maps are provided. A key is given. The mitochondrial gene cytochrome c oxidase subunit 1 (COI) and the nuclear gene 28S ribosomal RNA were sequenced for both species. COI di- vergence between some specimens of P. suterii was unexpectedly high (8.6%, uncorrected distance), but there was no morphological differentiation or 28S sequence divergence between those specimens. A molecular phylogenetic analy- sis examining the relationships between eight specimens of P. suterii and three specimens of P. keani using COI data is presented.

Keywords. Arachnida, Araneae, Periegopidae, New Zealand, Haplogynae, classification, distribution, ecology, biology, new species, key, phylogeny.

Vink, C. J.; Dupérré, N.; Malumbres-Olarte, J. 2013. Periegopidae (Arachnida: Araneae). Fauna of New Zealand 70: 41 pp.

Received: 5 July 2012. Accepted: 19 November 2012.

CHECKLIST OF TAXA Genus Periegops Simon, 1893...... 15 suterii (Urquhart, 1892)...... 15 keani new species...... 17

CONTENTS Phylogenetic analysis ...... 13 Acknowledgments ...... 8 Methods ...... 13 Introduction ...... 8 Results ...... 13 Morphology and terminology ...... 8 Discussion ...... 14 Methods and conventions ...... 10 Key to Periegopidae known from New Zealand ...... 15 Collecting ...... 10 Biosystematics ...... 15 Preservation ...... 10 References ...... 19 Preparation ...... 10 Appendices Measurements ...... 10 A: Glossary of technical terms ...... 23 Descriptions ...... 10 B: Collection details of specimens examined ...... 24 Illustrations ...... 11 Illustrations ...... 25 Text conventions ...... 11 Distribution maps ...... 34 Molecular biology ...... 11 Taxonomic index ...... 35 8 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

ACKNOWLEDGMENTS INTRODUCTION We are indebted to Thomas Buckley (New Zealand Arthro- Periegopidae Simon, 1893 is a family of six-eyed spiders pod Collection, Auckland) for his support of this project. comprising only three species, all in the genus Periegops We thank the following for the loan of specimens in Simon, 1893. Two species are endemic to New Zealand their care; Phil Sirvid (Museum of New Zealand Te Papa and the other is endemic to Queensland, Australia (Forster Tongarewa, Wellington), Grace Hall (New Zealand Ar- 1995). Periegopids can be distinguished from other fami- thropod Collection, Auckland), John Marris (Entomolo- lies found in New Zealand by a combination of characters. gy Research Museum, Lincoln University), Cody Fraser The six eyes are arranged in three, widely spaced diads. (Otago Museum, Dunedin), Robert Raven (Queensland The chelicerae have a lamina on the ventral surface and Museum, Brisbane), and Norman Platnick (American the maxillae are slender and are more than twice as long Museum of Natural History, New York). Thanks to as wide and directed across the labium. The anterior two Christine Rollard (Muséum national d’Histoire naturelle, pairs of legs have asymmetrical superior claws; the pro- Paris) for checking for the presence of the type of Per- claw of legs I and II has a double row of teeth whereas iegops hirsutus. the retroclaw has a single row (Forster 1995: fig. 11–13). Mike Bowie (Lincoln University) kindly supplied There is also a field of spicules on the median surface of specimens that have been valuable in this study. Mike the posterior median spinnerets (Labarque & Ramírez Fitzgerald and Grace Hall shared information on the 2012; fig. 21F, 22E) and the venom outlet is anterior on locations where Periegops specimens had been col- the fang (Labarque & Ramírez 2012; fig. 18A); both of lected. Thanks to John Kean (AgResearch) for collect- these characters require high microscope magnification ing assistance on Ruamahuanui Island, to Rob Chappell to see them. (Department of Conservation) for logistics support and Periegops suterii (Urquhart, 1892) was first de- to Kristine Grayson (Victoria University of Wellington) scribed from a female and two immature males collected for sharing her charter to the Aldermen Islands. Henry by the early New Zealand zoologist, Henry Suter, at Dy- Hudson Vink and Lucy Hudson Vink helped collect er’s Pass on the Port Hills near Christchurch (Urquhart specimens at Kennedys Bush Reserve. We thank Bryce 1892; Forster 1995). Urquhart placed his new species McQuillan for his excellent photographs of Periegops in a Northern Hemisphere genus and family; Segestria suterii. Latreille, 1804, and Dysderidae C.L. Koch, 1837, re- Facundo Labarque (California Academy of Scienc- spectively. Shortly after, Simon (1893) erected the genus es), Phil Sirvid, and Thomas Buckley provided helpful Periegops for his new species P. hirsutus, which he de- comments on an earlier draft of the manuscript. scribed from a female specimen sent to him at the Mu- Collecting permits were supplied by the New Zea- séum national d’Histoire naturelle, Paris, from New Zea- land Department of Conservation (permits WK-30010- land. Although Simon (1893) did not specify the location FAU and CA-28815-FAU). The Ruamaahua Island Trust where the specimen was collected from, it was likely that granted cultural permission to collect on Ruamahuanui it was from Banks Peninsula (Forster 1995). The gener- Island. This research was supported by Core funding for ic description was of a female and Simon (1893: 267) Crown Research Institutes from the Ministry of Busi- noted that the male was unknown; however, the descrip- ness, Innovation and Employment’s Science and Inno- tion of P. hirsutus was listed as male although no male vation Group through the Defining New Zealand’s Land characteristics appear to be described (Simon 1893: 268). Biota Portfolio. Bryant (1935a: 54) and Forster (1995: 92) noted that the listing as a male appears to be in error. Dalmas (1917: 338) also noted the specimen was female and this was the specimen seen by Chamberlain (1946). It is unlikely that Simon realised he had described the same species as Urquhart and P. hirsutus was later made a synonym of P. suterii by Chamberlain (1946). Simon (1893) was first to recognise that the genus Periegops was distinct from other genera and placed it in its own subfamily, Periegopinae Simon, 1893. Forster (1995) elevated the subfamily to family status when he described Periegops australia Forster, 1995 and redescribed P. suterii. For- ster (1995: 96) also noted that he had examined a single Fauna of New Zealand 70 ­9­ female specimen collected at East Cape, but there were of the Haplogynae. Periegopidae was not included in the “no clear characters by which the species could be satis- analyses but Ramírez (2000) stated that Periegopidae factorily defined”; a male would need to be collected and was sister to the Scytodidae. Periegopidae was also in- the palpal bulb examined to determine if it was a sepa- cluded in the scytodoid clade by Coddington (2005); its rate species. Forster & Forster (1999) mentioned an ad- inclusion was presumably based on Forster (1995) and ditional specimen from the Aldermen Islands, which they Ramírez (2000). Labarque & Ramírez (2007a, b) consid- believed to be a female, and Vink (2006) reported that he ered that among the scytodoids, Periegopidae was closest had sequence data from a fragment of the mitochondrial to Drymusidae and Scytodidae as they share two char- gene cytochrome c oxidase subunit 1 (COI) [specimen acters; asymmetrical superior claws on the anterior two Pk1] that suggested that the North Island specimens were pairs of legs and a field of spicules on the median surface a distinct species. of the posterior median spinnerets. More recently, La- Periegopidae is part of a group of spiders called the barque & Ramírez (2012) expanded the morphological Haplogynae Simon, 1893, which are araneomorph spi- dataset of Platnick et al. (1991) and Ramírez (2000) to ders that lack separate fertilisation ducts, do not have a include Periegopidae along with a restricted Scytodoi- sclerotised epigyne and the male pedipalp is relatively dea, which included Drymusidae, Scytodidae, and Si- simple. Haplogynae are a monophyletic group (Plat- cariidae. Periegops, Drymusa, and Scytodidae formed a nick et al. 1991; Ramírez 2000) that includes 17 fami- clade, which all shared the synapomorphies of asymmet- lies. Perigopids have a number of characteristics that rical superior claws and a field of spicules on the median are shared with the haplogyne families Diguetidae F. O. surface of the posterior median spinnerets. Labarque & Pickard-Cambridge, 1899, Drymusidae Simon, 1893, Ramírez (2012) concluded that Periegopidae was sister Plectreuridae Simon, 1893, Scytodidae Blackwall, 1864, to Drymusidae, but they did not find any autapomorphies and Sicariidae Keyserling, 1880. These include a lamina for Periegops and suggested that the two families might on the ventral surface of the chelicerae, slender maxillary be synonymised. lobes directed across the labium, and a limited posterior Very little is known of the biology of periegopids. respiratory system with fused apodemes (Forster 1995). They are only found in forest with a deep leaf litter layer Forster (1995) grouped this set of families, along with and well-drained soil. Specimens have been collected Periegopidae, into the superfamily Sicarioidea, but did from under logs and rocks, in leaf litter, in pitfall traps, not provide a formal definition of the superfamily. The in rotted logs and root cavities, and in a mygalomorph name Sicarioidea was first used by Berland (1932) and burrow (Forster 1995; Vink 2006). Forster (1995) sug- Forster (1995) appears to be the only other arachnologist gested that periegopids were rare as their habitat has been that has used it. severely reduced due to human deforestation and only A more commonly used superfamily name that in- 21 Periegops specimens had been collected in New Zea- cludes this set of taxa is Scytodoidea (Bristowe 1938; land and Australia between 1891 and 1995. In surveys Caporiacco 1938; Brignoli 1978; Lehtinen 1986); how- between October 2002 and May 2003, Vink (2006) found ever, Scytodoidea has been used to refer to a larger set a further 24 specimens of P. suterii at ten different loca- of haplogyne families that has also included Caponiidae tions on the Banks Peninsula. Sirvid et al. (2012) classi- Simon, 1890, Leptonetidae Simon, 1890, Ochyrocerati- fied P. suterii as having a relict distribution. dae Fage, 1912, Pholcidae C. L. Koch, 1850, and Tetrab- Periegops do not appear to build a web for prey cap- lemmidae O. Pickard-Cambridge, 1873. A morphological ture, but do build silken retreats. It seems probable that phylogenetic analysis by Platnick et al. (1991) supports periegopids are cursorial, night-time hunters as they have a clade informally named "scytodoids" (Coddington & only been found on the forest floor, have been collected Levi 1991) that includes Diguetidae, Drymusidae, Lep- in pitfall traps and lack webs. On two occasions a single tonetidae, Ochyroceratidae, Pholcidae, Plectreuridae, female has been found together with two or three males Scytodidae, Sicariidae, Telemidae Fage, 1913, and Tet- under logs and rocks (Forster 1995; Vink 2006), which rablemmidae. The monophyly of this clade is supported implies that the female might possess some method of by the reduction of the posterior spiracles to one and the attracting males. tetrahedral posterior median spinnerets (Platnick et al. 1991), although these characters have been reversed in some species in the clade (Platnick et al. 1991). MORPHOLOGY AND TERMINOLOGY Ramírez (2000) examined the morphology of haplo- gyne respiratory systems and added characters to the data The morphological structures required for the identifica- matrix of Platnick et al. (1991) to produce a phylogeny tion of New Zealand and Australian Periegopidae are 10 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae) referred to in Fig. 1–16 and explained in the glossary of Preparation. Specimens should be labelled with the technical terms (Appendix A), and Paquin et al. (2010). locality, including area code (Crosby et al. 1976, 1998), As with almost all spiders, the male pedipalp is important latitude and longitude, collection date, collector’s name, when identifying periegopids to species. The female epi- habitat data, and collection method. gyne, which is usually useful for identifying spider species Most morphological features used in identification (Paquin et al. 2010), is just a lightly sclerotised area that can be seen under an ordinary dissecting microscope. covers the internal genitalia in periegopids, and many When examining a spider in ethanol it should be rested other haplogyne spiders, and therefore is not likely to be in washed quartz sand or glass beads to provide support diagnostic. The internal genitalia, however, are diagnostic, for the specimen. This also allows the specimen to be as is the case with most spider species. The morphological positioned at any desired viewing angle. The features nomenclature of the pedipalp and the internal genitalia of the male pedipalp are best viewed by removing the follows Forster (1995). left pedipalp at the junction between the trochanter and A phylogenetic species concept (Wheeler & Platnick the femur, and viewed ventrally. In periegopids, as with 2000) has been implemented in this study. It defines a other haplogyne spiders, there is no sclerotised external species as the smallest aggregation of populations diag- genitalia, only a slightly sclerotised plate, which makes nosable by a unique combination of character states. distinguishing females from immature specimens diffi- cult. Female internal genitalia were excised using a sharp entomological needle and were prepared for examination by placing the dissected genitalia in either lactic acid or 10% KOH solution for 1–3 hours at 37°C to dissolve soft METHODS AND CONVENTIONS tissue. Internal genitalia were only illustrated for P. sute- Collecting. Periegopids can be collected by a variety of rii and P. australia. methods. The best method for collecting periegopids is by searching under rocks and logs that are sitting on or Measurements. All measurements are in millimetres partially embedded in soil on open ground within forest in (mm). Where the measurements are expressed as a frac- areas with a good litter layer over well drained soil (Vink tion, the numerator refers to the length of the structure and 2006). Periegopids have also been found under wooden the denominator refers to its width. Measurements outside discs (M. H. Bowie, pers. comm.) that are facsimiles for parentheses are for males and inside parentheses for fe- natural fallen logs (Bowie & Frampton 2004). Periegopids males. The size ranges given for the body and carapace have also been collected by sieving and searching leaf length of each species represent the smallest and largest litter. Specimens of P. suterii and P. australia have been individuals of each sex that we examined. The mean body caught in pitfall traps, but unless specimens are collected length and carapace length were calculated and the number and preserved within a couple of days of being caught they of specimens measured given. can start to decay, which can make identification difficult. Decay of specimens caught in pitfall traps can be mitigated Descriptions. For the new species, illustrations, measure- by the use of a good preservative such as propylene gly- ments and colour pattern descriptions were made from col, which also preserves DNA but can shrivel soft tissue the type specimen. For the existing species, illustrations, (Vink et al. 2005). Malumbres-Olarte et al. (in press) used measurements, and colour pattern descriptions were propylene glycol in their pitfall traps and found that useful prepared from a non-type representative male and female DNA was preserved after two weeks in the field. specimen (with collection information shown). Descriptions of colours are for ethanol-preserved Preservation. Periegopids are best preserved long-term specimens. It should be noted that colours and colour in 70–75% ethanol. To ensure adequate DNA preserva- patterns can fade in older specimens, particularly those tion, specimens should also be stored at ≤ -20°C (Vink et that have not been stored away from light. al. 2005). Spiders can be stored in 95–100% ethanol to Measurements were made with a micrometer ruler preserve DNA but as with lower ethanol concentrations, fitted to the eyepiece of a Leica M125 stereo microscope. it is still best to combine this with storage temperatures ≤ Characters diagnostic in other spider families (e.g., -20°C (Vink et al. 2005). Preservation in 95–100% etha- eye size and position, leg spination) are not diagnostic nol makes specimens brittle, potentially rendering them for Periegopidae species and have not been included in unsuitable for morphological examination. the descriptions. Fauna of New Zealand 70 ­11­

Illustrations. Specimens to be illustrated were first CAACAAATCATAAAGATATTGG-3’) (Folmer et photographed with a Nikon Coolpix 950 digital camera al. 1994) plus C1-N-2776-spider (5’-GGATAAT- attached to a SMZ-U Nikon dissection microscope. The CAGAATANCGNCGAGG-3’) (Vink et al. 2005) or digital photos were then used to establish proportions and C1-J-1718-spider (5’-GGNGGATTTGGAAATTGRT- the illustrations were detailed and shaded by referring back TRGTTCC-3’) (Vink et al. 2005) plus C1-N-2776-spi- to the structure under the microscope. der. The two COI fragments were 1260 base pairs (bp) Map images were created using the geographic infor- and 1055 bp long, respectively. Two different primer mation system software ArcMap 10 (ESRI). pairs were used to amplify and sequence two over- lapping 28S fragments; 28S-B1 (5'-GACCGATAG- Text conventions. The area codes of Crosby et al. (1976, CAAACAAGTACCG-3') (Bruvo-Mađaric et al. 2005) 1998) are used in collection records. plus 28S-B2 (5'-GATTAGTCTTTCGCCCCTATA-3') The following acronyms for repositories are used: (Bruvo-Mađaric et al. 2005) and 28Sa (5'-GACCC- AMNH American Museum of Natural History, New GTCTTGAAACACGGA-3') (Nunn et al. 1996) plus York, U.S.A. LSUR (5'-GCTACTACCACCAAGATCTGCA-3') (Rix LUNZ Entomology Research Museum, Lincoln et al. 2008). The two 28S fragments were 819–821 bp University, New Zealand and 830–833 bp long, respectively. PCR amplification MONZ Museum of New Zealand Te Papa Tongarewa, was performed using i-StarTaq™ DNA Polymerase Wellington, New Zealand (iNtRON Biotechnology) in a Mastercycler® (Eppen- NZAC New Zealand Arthropod Collection, Auckland, dorf) thermocycler with a cycling profile of 35 cycles New Zealand of 94ºC denaturation (30 s), 48°C (COI) or 60°C (28S) OMNZ Otago Museum, Dunedin, New Zealand annealing (30 s), 72ºC extension (1 min) with an initial QMB Queensland Museum, Brisbane, Australia denaturation of 3 min and a final extension of 5 min. Ex- cess primers and salts were removed from the resulting double-stranded DNA using a DNA Clean & Concen- Molecular biology. To construct a molecular phylogeny of trator™ Kit (Zymo Research). Double bands were ob- New Zealand Periegopidae and to facilitate the identifica- served when 28S PCR products of Pk1 were visualised tion of immature specimens, we used the mitochondrial via gel electrophoresis; both bands were excised from the gene cytochrome c oxidase subunit 1 (COI) and the nuclear gel and prepared for sequencing using a Zymoclean™ ribosomal RNA gene 28S. COI is one of the fastest evolv- Gel DNA Recovery kit. For Pk1, the primer pair 28S- ing mtDNA genes and has been used to examine genetic B1 plus 28S-B2 amplified a fragment of 28S DNA that differences between and among haplogyne spiders (e.g., BLAST database searching indicated may have come Arnedo et al. 2001, 2009; Astrin et al. 2006; Starrett & from a ciliate protozoan. For Pk2, the primer pair 28Sa Waters 2007; Binford et al. 2008; Dimitrov et al. 2008; plus LSUR amplified a fragment of 28S DNA from a soil Huber & Astrin 2009; Duncan et al. 2010) and New Zea- nematode in the family Cephalobidae. Amplification of land spider species (Vink & Paterson 2003; Vink et al. 28S was attempted for specimens Ps5 and Ps6, but was 2008, 2011a, b; Framenau et al. 2010; Vink & Dupérré not possible owing to contamination by a fungus that the 2010; Lattimore et al. 2011; Malumbres-Olarte & Vink primers preferentially annealed to. Contaminants were 2012). 28S was selected because it is a slow evolving identified by BLAST searching their sequences (Altschul gene (Hedin & Maddison 2001) and has been used in et al. 1997). Purified PCR fragments were sequenced phylogenetic analyses of haplogynes (Bruvo-Mađaric in both directions at the Core Instrumentation Facility et al. 2005; Binford et al. 2008) as well as other spiders (University of California, Riverside, USA), the Massey (e.g., Hedin & Bond 2006; Rix et al. 2008; Wang et al. Genome Service (Massey University, New Zealand), 2008; Spagna et al. 2010). Eight specimens of P. suterii or Macrogen (Korea). Sequence data were deposited in and three specimens of P. keani were sequenced for COI. GenBank (www.ncbi.nlm.nih.gov/GenBank/ – see Table Each specimen was arbitrarily assigned a specimen code 1 for accession numbers). Sequences were edited using (Table 1). A subset of four specimens, two from each spe- Sequencher 4.6 (Gene Codes Corporation). cies, was sequenced for 28S. DNA was extracted non-destructively (see Pa- quin & Vink 2009) from either two to three legs us- ing a ZR Genomic DNA™ Tissue Minipreps (Zymo Research). The primers used to PCR amplify and se- quence COI fragments were either LCO1490 (5’-GGT- 12 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae) GenBank accession numbers JX174286, JX174291 JX174287 JX174288, JX174292 JX174289, JX174293 JX174281 JX174282 JX174283 JX017357, JX017353 JX174284 JX174285 JX174290 Collection information MC, Montgomery Park Reserve, 43°44.7’S, 172°52.2’E, LUNZ (00012725) 5 November 2010, JM-O & CJV, MC, Kennedys Bush Reserve, 43°37.9’S, 172°37.3’E, LUNZ Hudson Vink, CJV & H.P. August 2011, 13 (00012718) CL, Ruamahuanui Island, 36°57.25’S, 176°05.52’E, 19 November 2003, B.M. Fitzgerald, MONZ (AS.002310) CL, Ruamahuanui Island, 36°57.25’S, 176°05.52’E, 23 March 2012, CJV & J.M. Kean, LUNZ (00012716) MC, Hinewai Reserve, 43°48.59’S, 173°01.28’E, 11 MC, Hinewai Reserve, 43°48.59’S, 173°01.28’E, 11 AMNH (ARAMR-000615) October 2002, CJV, MC, Hinewai Reserve, 43°48.59’S, 173°01.28’E, 11 AMNH (ARAMR-000616) October 2002, CJV, MC, Hay Reserve, 43°42.18’S, 172°53.87’E, 13 March LUNZ (00012720) 2003, CJV, MC, Kennedys Bush Reserve, 43°37.9’S, 172°37.3’E, LUNZ & L.J. Hudson Vink, H.P. CJV, April 2011, 22 (00012715) MC, Montgomery Park Reserve, 43°44.7’S, 172°52.2’E, 5 November 2010, CJV & JM-O, LUNZ (00012722) MC, Hinewai Reserve, 43°48.69’S, 173°00.95’E, 11 M.H. Bowie, LUNZ (00012735) January 2011, CL, Ruamahuanui Island, 36°57.25’S, 176°05.52’E, 23 March 2012, CJV & J.M. Kean, LUNZ (00012717) ♀ ♂ ♂ ♀ ♀ ♂ ♂ ♂ Sex immature immature subadult ♀ Ps8 Pk1 Pk2 Pk3 Ps1 Ps2 Ps3 Ps4 Ps5 Ps6 Ps7 code Specimen Species Periegops suterii Periegops keani Periegops keani Periegops suterii Periegops suterii Periegops suterii Periegops suterii Periegops suterii Periegops suterii Periegops suterii Periegops keani Table 1 . Specimens used for molecular analysis. Table Fauna of New Zealand 70 ­13­

Table 2. Uncorrected distance matrix for cytochrome c oxidase subunit 1 (COI).

Pk1 Pk2 Pk3 Ps1 Ps2 Ps3 Ps4 Ps5 Ps6 Ps7 Pk2 0.006 Pk3 0.007 0.007 Ps1 0.135 0.131 0.135 Ps2 0.135 0.131 0.135 0.000 Ps3 0.139 0.135 0.139 0.007 0.007 Ps4 0.141 0.138 0.138 0.084 0.084 0.086 Ps5 0.132 0.130 0.132 0.012 0.012 0.013 0.082 Ps6 0.135 0.131 0.135 0.000 0.000 0.007 0.084 0.012 Ps7 0.132 0.130 0.132 0.012 0.012 0.013 0.082 0.000 0.012 Ps8 0.141 0.138 0.138 0.084 0.084 0.086 0.000 0.082 0.084 0.082

PHYLOGENETIC ANALYSIS deviation of split frequencies had dropped below 0.002, which indicated that the two tree samples had converged. Methods. Sequences were aligned using Sequencher Tracer version 1.5 (Rambaut & Drummond 2009) was 4.6. There was no evidence of insertions/deletions or also used to determine if the analyses had sufficient ef- stop codons in the COI sequences and alignment was fective sample sizes. MrBayes was used to construct ma- straightforward. jority rule consensus trees, discarding the first 25% of Uncorrected COI pairwise distances were calculated trees generated as burn-in. TreeView 1.6.6 (Page 1996) using PAUP* version 4.0b10 (Swofford 2002). Distances was used to view and save trees in graphic format. using the Kimura-2-Parameter (K2P) model (Kimura 1980) were also calculated for comparison with previ- Results. A 1031 base pair (bp) COI fragment was se- ously reported intraspecific distances in spiders (Robin- quenced from eight specimens of P. suterii and three son et al. 2009). Although K2P has been commonly used specimens of P. keani. Four COI haplotypes occurred in most DNA barcoding studies (e.g., Robinson et al. among the eight specimens of P. suterii. Each of the three 2009), there is no evidence that this model is better for specimens of P. keani had a different COI haplotype. Inter- species identification than simpler metrics such as uncor- and intraspecific uncorrected pairwise distances between rected pairwise distances (Astrin et al. 2006; Collins et COI sequences of P. suterii and P. keani are shown in Table al. 2012). 2. The minimum divergence between the two species was Partitioned Bayesian analysis implemented in 13.0% (uncorrected) and the mean divergence was 13.5% MrBayes version 3.1.2 (Ronquist & Huelsenbeck 2003) (uncorrected). The mean intraspecific divergences in P. was used to estimate the COI phylogenetic tree topology. suterii and P. keani, were 4.0% (uncorrected) and 0.6% MrModeltest version 2.3 (Nylander 2008) implemented (uncorrected), respectively. The maximum intraspecific in PAUP* version 4.0b10 (Swofford 2002) was used to divergence in P. suterii was 8.6% (uncorrected, 9.2% K2P- select the optimal model and model parameters. Within distance) and was 0.7% (uncorrected and K2P-distance) MrModeltest the Akaike Information Criterion was used in P. keani. for model selection (Posada & Buckley 2004). Based on The 1360 bp fragments of 28S from two specimens the results of Brandley et al. (2005), the COI data were of P. suterii were identical. The amplification of 28S partitioned by codon with models selected for each co- from two specimens of P. keani (Pk1 and Pk2) was only don; GTR+I (Lanave et al. 1984; Tavaré 1986) for the partially successful with an 821 bp fragment from Pk1 st rd 1 and 3 codon positions and HKY+I (Hasegawa et and an 833 bp fragment from Pk2; however, the 281 bp nd al. 1985) for the 2 codon positions. Bayesian analysis overlap was identical. There were 12 nucleotides that was conducted by running two simultaneous, complete- varied between the two species. ly independent analyses each with four heated chains, The phylogenetic analysis of the COI data (Text-fig. th sampling every 1000 tree. The analysis was run for 20 1) showed that P. suterii and P. keani are monophyletic million generations, by which time the average standard with P. suterii divided into two clades. 14 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Discussion. Periegops COI sequences were very similar or identical (Table 2) for specimens collected at the same sites; however, specimens of P. suterii collected at Kenne- dys Bush Reserve, in the west of the Banks Peninsula (Map 1), were 8.2–8.6% (8.7–9.2% K2P-distance) divergent from P. suterii specimens collected from three localities in the eastern Bank Peninsula (Map 1). This high divergence was not found in the slower evolving nuclear marker; 28S sequences were identical from specimens from Kennedys Bush Reserve and Montgomery Park Reserve (Map 1). We also did not observe any morphological differences be- tween Kennedys Bush Reserve specimens and specimens collected from other locations on the Banks Peninsula; therefore we are confident that all specimens are one spe- cies, P. suterii. A difference of 8.7–9.2% (K2P-distance) in COI sequence divergence is very high in spiders; much more than the average maximum intraspecific divergence of 3.16% (K2P-distance) observed in other spiders (Robin- son et al. 2009). However, higher intraspecific divergences (>10%) have been observed in other haplogynes (Astrin et al. 2006; Binford et al. 2008), Hypochilidae (Hedin 2001), and Mygalomorphae (Bond et al. 2001), so perhaps COI divergence is higher in ancestral spiders than it is in the En- telegynae. Nevertheless, it is curious that high (8.2–8.6%) Text-fig. 1 Unrooted Bayesian consensus tree based COI divergence was observed between specimens from lo- on cytochrome c oxidase subunit I (COI) sequence calities only just over 23 km apart and at similar altitudes, data. Values on branches are posterior probabilities. while there was much less divergence (0.7–1.3%) between Specimen codes are listed in Table 1. Branch specimens from eastern Bank Peninsula localities, which lengths are proportional to the expected number of were 5–15 km apart. Specimens from localities between substitutions per site (see scale bar). Kennedys Bush Reserve and Montgomery Park Reserve, such as Kaituna Valley, could reveal if COI divergence is correlated with distance or whether there has been some sort of genetic isolation between the east and west of Banks Peninsula, similar to the geographic distribution of two closely related weta species in the genus Hemideina (Townsend et al. 1997). Fauna of New Zealand 70 ­15­

KEY TO NEW ZEALAND PERIEGOPIDAE of leg I with a row of strong setae on retrolateral surface (Fig. 4, 5). Proclaw of legs I and II with double row of 1 Tibia of leg I less than 0.7× length of carapace. When teeth, retroclaw with single row. Both superior claws of viewed retrolaterally, dorsal surface of cymbium of legs II and IV a single row. Inferior claw of all tarsi with a male pedipalp projects dorsally (Fig. 9). Flared flat- single tooth. Male pedipalp simple with a bulb and a short tened flanges near embolus tip (Fig. 6, 7). Found only embolus (Fig. 6–9); coiled sperm duct often visible within on Banks Peninsula and Riccarton Bush (Map 1). .... the bulb. Abdomen ovoid, dun or dirty cream usually with ...... (p. 15)... Periegops suterii (Urquhart) a distinct chevron pattern (Fig. 1, 14–16) and black setae. ––Tibia of leg I more than 0.75× length of carapace. Slightly sclerotised area anterior to the epigastric furrow When viewed retrolaterally, dorsal surface of cym- (Fig. 10, 11). Female internal genitalia simple, haplogyne, bium of male only slightly projects dorsally (Fig. 9). consisting of a rounded poreplate with many small invagi- Flattened flanges near embolus tip not flared (Fig. nated cups (Fig. 12, 13). 6 spinnerets and small colulus; 9a). Found only on Aldermen Islands and East Cape median surface of posterior median spinnerets with field (Map 2)...... (p. 17)…Periegops keani sp. nov. of spicules (Labarque & Ramírez 2012; fig. 21F, 22E). Forster (1995) provided details of the respiratory system of Periegops.

BIOSYSTEMATICS Periegops suterii (Urquhart) Fig. 1, 4, 6, 7, 10, 12, 14, 16; Map 1 Family PERIEGOPIDAE Segestria suterii Urquhart, 1892: 230. Periegops hirsutus Simon, 1893: 268, figs 223–224.—Synonymy Medium-sized, 3-clawed haplogyne spiders. 6 eyes in 3 by Chamberlain 1946: 88. widely separated diads (Fig. 1, 14, 15). Proclaw of legs Periegops suteri (Urquhart) —Bryant 1935a: 53. —Bryant 1935b: I and II with double row of teeth, retroclaw with single 81, figs 4, 13, 25. —Chamberlain 1946: 88. —Forster 1967: row. Chelicerae with lamina on the ventral surface (Fig. 72, figs 81, 134. —Forster & Forster 1973: 151, figs 87b, 90. 2) and slender maxillary lobes directed across the labium —Jocqué & Dippenaar-Schoeman 2006: 202, figs 78a–e. —Labarque & Ramírez 2012: 1, figs 1, 2A, 3A–B, 6A–B, (Fig. 3). Male pedipalp simple, consisting of a bulb with a 9A, 10A, 11A, 12A, 13A, 14A, 15A, 16A, 17A, 20A, 21, 22, short embolus (Fig. 6–9). Slightly sclerotised area anterior 29A–B, 31A. to the epigastric furrow (Fig. 10, 11) and simple internal Periegops suterii (Urquhart) —Forster 1995: 94, figs 1–3, 6–14, genitalia (Fig. 12, 13). 23, 24, 27, 31. —Forster & Forster 1999: 143, figs 1.14i, 10.9. —Paquin, Vink & Dupérré 2010: 37, figs 14.1–4. Diagnosis. Distinguished from all other Periegops species Genus Periegops Simon by the shape of the embolus and cymbium of the male pedi- Periegops Simon, 1893: 267. —Forster 1995: 93–94. Type species palp. Flattened flanges near embolus tip more pronounced Periegops hirsutus Simon, 1893. than those of P. keani, and embolus is shorter than that of Description. Body length 5.9–10.0 mm. Carapace longer P. australia. Dorsal surface of cymbium projects slightly than wide, flat, without fovea, orange or orange-brown, more than that of P. keani, but much less than that of P. darker anteriorly. 6 eyes in 3 widely separated diads; australia. Poreplate of internal genitalia not as rounded as anterior median eyes absent. Chelicerae with lamina on that of P. australia. Leg I of males, females, and immatures ventral surface and membranous promarginal lobe with shorter than in P. keani; tibia I less than 0.7× length of 5–7 setae (Fig. 2); 3 promarginal teeth and 1 reduced carapace. Found only on Banks Peninsula and Riccarton retromarginal tooth; fang length less than width of patu- Bush in the South Island. ron; venom outlet anterior on fang (Labarque & Ramírez Description. Carapace: red-orange anteriorly, orange pos- 2012; fig. 18A). Lateral surface of female chelicerae teriorly; with sparse black setae; fovea absent. Sternum: with stridulatory ridges (Labarque & Ramírez 2012; fig. orange. Leg I orange-brown, patella, distal ends of femur 16A), with associated palpal femoral thorns (Labarque & and tibia light orange; legs II, III, and IV yellow-brown, Ramírez 2012; fig. 11A). Maxillae slender and more than darker at proximal end of each leg segment. Abdomen: 2× as long as wide, directed across the labium (Fig. 3). dun with black setae; black-brown chevron pattern (Fig. Legs in descending order of length usually I, IV, II, III, but 1, 14, 16). Chelicerae red-brown. Male pedipalp simple legs I and IV sometimes subequal in females. Male tibiae with a bulb and a short embolus, which is flattened and 16 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae) twisted, flaring out to 2 flanges just before the tip (Fig. 6, primary types held in Canterbury Museum (Nicholls et al. 7); coiled sperm duct often visible within the bulb. Slightly 2000). It was not possible to check whether the type speci- sclerotised area anterior to epigastric furrow (Fig. 10). men was present at Canterbury Museum, as the collections Female internal genitalia simple, haplogyne, consisting have not been accessible since the Christchurch earthquake of a rounded poreplate with many small invaginated cups in February 2011 (S. D. Pollard, pers. comm.). The type is (Fig. 12). not in OMNZ (C. Fraser, pers. comm.), which houses New Dimensions (mm). Male MC, Kennedys Bush Reserve Zealand’s largest collection of spiders. It is possible that LUNZ 00012715 (female MC, Kennedys Bush Reserve the type was amongst specimens at Ray Forster’s house at LUNZ 00012718): total length 6.13 (7.92); carapace the time of his death and has as yet not been located. We 3.06/2.05 (3.82/2.60), height 1.15 (1.71); abdomen consider the type of Segestria suterii missing, but not lost. 2.87/1.80 (4.12/2.60); sternum 1.64/1.05 (1.93/1.25). Type of Periegops hirsutus: Not examined. Original description based on a female specimen from an undis- I II III IV Palp closed location in New Zealand (Simon 1893). Although many of Simon’s type specimens are housed in the Mu- 2.43 2.13 1.84 2.35 0.65 Femur séum national d’Histoire naturelle, Paris, France, the (2.72) (2.41) (2.1) (2.56) (0.95) type of Periegops hirsutus could not be found there (C. 0.85 0.8 0.73 0.8 0.27 Patella Rollard, pers. comm.). (0.98) (0.98) (0.9) (0.97) (0.38) Material examined. 32 non-type specimens (15 males, 2.08 1.75 1.45 2.03 0.6 Tibia (2.15) (1.93) (1.65) (2.3) (0.63) 15 females, 2 subadult females) — see Appendix B for collection details of specimens examined. 2.16 1.82 1.62 2.07 Metatarsus (2.17) (1.97) (1.78) (2.28) Distribution (Map 1). Found only on Banks Peninsula 0.88 0.76 0.68 0.68 0.22 and Riccarton Bush (MC). Forster (1995) and Vink (2006) Tarsus (0.85) (0.83) (0.73) (0.73) (0.78) listed localities where P. suterii specimens have been 8.4 7.26 6.32 7.93 1.74 found. Riccarton Bush is the only site outside of the Banks Total (8.87) (8.12) (7.16) (8.84) (2.74) Peninsula where P. suterii has been found (in 1994), but subsequent searches there (Chinn 2006; Vink 2006) have failed to find it. Variation. Size range, mean. Male body length 5.9–7.4, 6.6, n = 12. Female body length 6.8–9.3, 8.1, n = 11. Male Biology. Periegops suterii has been found in forest with carapace length 3.0–3.7, 3.4, n = 12. Female carapace a deep leaf litter layer and well-drained soil. It has been length 3.4–4.8, 3.9, n = 11. found under logs and rocks, and in leaf litter, in both beech The ratios of the length of tibia I to the length of the and podocarp forest. Within suitable locations, P. suterii carapace (measured for all specimens, including imma- has a patchy distribution, but specimens are often found tures) were between 0.54 and 0.70. The ratio of leg I seg- years apart in the same specific locations.Periegops suterii ment length to carapace length has also been shown to does not appear to build a web for prey capture, but it does be a reliable character for the separation of other closely produce dragline silk and builds silken retreats. The lack related spider species (Vink et al. 2008). of webs and their presence in pitfall traps suggest that it is The chevron pattern on the abdomen (Fig. 1, 14, 16) a cursorial, night-time hunter. On two occasions a single was visible in all but one of the specimens examined, a female has been found together with two or three males male (LUNZ 00012715). Forster (1995) noted that a fe- under logs and rocks (Forster 1995; Vink 2006), which male collected at Riccarton Bush did not have abdominal implies that the female might possess some method of at- markings. tracting males. Forster (1995) suggested pheromones, but it could also be stridulation; Forster (1995) did not notice Type data. Type of Segestria suterii: Not examined. Origi- the stridulatory ridges on the chelicerae. Adults have been nal description based on a female specimen from “Dyer’s found from August to May. Pass, Canterbury”, which was collected with 2 immature males by H. Suter (Urquhart 1892). Forster (1995) stated DNA. Nine cytochrome c oxidase subunit 1 and two 28S the type was in poor condition in the Canterbury Museum, rRNA sequences for this species are listed in GenBank Christchurch, New Zealand, where all of Urquhart’s ex- (www.ncbi.nlm.nih.gov/GenBank/) under accession isting type specimens are housed (Court & Forster 1988; numbers EF537066, JX017353, JX017357, JX174281– Nicholls et al. 2000; Paquin et al. 2008). However, the type JX174287, and JX174291. of Segestria suterii was not included in the list of arachnid Fauna of New Zealand 70 ­17­

Remarks. Although the types of Segestria suterii and orange-brown, patella, distal ends of femur, and tibia Periegops hirsutus were not examined, we have no reason orange-brown; legs II, III, and IV orange-brown, darker at to doubt that these species are synonymous, as Chamber- proximal end of each leg segment. Abdomen: dirty cream lain (1946) examined the type specimens of both species. with black setae; chevron pattern barely visible (Fig. 15). Forster (1995: 92) stated that it can “be reliably assumed” Chelicerae red-brown. Male pedipalp simple with a bulb that the type of P. hirsutus “came from the same locality and a short embolus, which is flattened and twisted, flaring as Urquhart’s Segestria suterii.” Also, Simon (1893) noted out slightly to 2 flanges just before the tip (Fig. 9); coiled that the type specimen of P. hirsutus was 8 mm long, which sperm duct visible within the bulb. Female unknown. is within the range of female body lengths we observed Dimensions (mm). Male holotype: total length 10.03; for P. suterii. carapace 4.99/3.48, height 1.66; abdomen 5.25/3.75; Despite the common usage of the name Periegops sternum 2.40/1.90. suteri, the correct original spelling of the specific epithet is suterii (as Segestria suterii Urquhart, 1892). Article I II III IV Palp 33.4 in the fourth edition of the International Code of Zoological Nomenclature (International Commission on Femur 4.99 4.26 3.73 4.41 1.01 Zoological Nomenclature 1999) states the change from Patella 1.41 1.08 1.21 1.26 0.45 -i to -ii is an incorrect subsequent spelling. There is pro- vision in the International Code of Zoological Nomen- Tibia 4.39 3.83 2.97 4.13 0.96 clature for prevailing usage of an unjustified emendation Metatarsus 4.44 3.88 3.23 3.98 (article 33.2.3.1) or incorrect subsequent spelling (article 33.3.1); however, suteri is not an emendation but an in- Tarsus 1.31 1.26 1.01 1.26* 0.45 correct subsequent spelling and article 33.4 is an excep- tion to articles 33.2.3.1 and 33.3.1. Both spellings have Total 16.54 14.31 12.15 15.04 2.87 been used; P. suterii by Forster(1995), Forster & Forster (1999), Paquin et al. (2010), and Sirvid et al. (2011, *Both tarsi IV were damaged so the length was estimated 2012), and P. suteri by Bryant (1935a, b), Chamberlain based on the proportions observed in immature species (1946), Forster (1967), Forster & Forster (1973), Vink of P. keani. (2006), Jocqué & Dippenaar-Schoeman (2006), Brock- Variation. The holotype male was the only specimen that erhoff et al. (2008), Platnick (2012), and Labarque & did not have abdominal markings; all of the other subadult Ramírez (2012). As this is not a case of prevailing usage and immature specimens had a chevron pattern on the of an incorrect subsequent spelling (article 33.3.1) and abdomen, like that of P. suterii (Fig. 1, 14, 16). article 33.4 is quite clear on the matter, we advocate for The ratios of the length of tibia I to the length of the and use the correct original spelling. carapace (measured for all specimens, including imma- tures) were between 0.77 and 0.88. Based on what we ob- served for P. suterii, it seems likely that female P. keani Periegops keani new species will have a tibia/carapace ratio within that range. Fig. 2, 3, 5, 9, 15; Map 2 Type data. Holotype: male (LUNZ 00012716) labelled “NEW ZEALAND, CL, Ruamahuanui Island; 36°57.25'S, Diagnosis. Distinguished from all other Periegops species 176°05.52'E; NW campsite, under rock; 23 Mar 2012, C.J. by shape of embolus and cymbium of the male pedipalp. Vink & J.M. Kean” (LUNZ). Flattened flanges near embolus tip not as pronounced as those of P. suterii and embolus shorter than that of Material examined. Type specimen plus 7 non-type P. australia. Dorsal surface of cymbium projects only specimens (1 subadult male, 1 subadult female, 5 im- slightly, less than that of P. suterii and much less than that matures) — see Appendix B for collection details of of P. australia. Leg I of male and immatures longer than specimens examined. in P. suterii; tibia I more than 0.75× length of carapace. Distribution (Map 2). Found only on the Aldermen Islands Found only on the Aldermen Islands and East Cape in and East Cape (CL, GB). the North Island. Biology. Periegops keani has been found in forest with a Description. Carapace: dark orange-brown anteriorly, deep leaf litter layer and well-drained soil. On the forest orange-brown posteriorly; with sparse black setae; fo- floor it has been found under rocks and in aStanwellia sp. vea absent. Sternum: light orange brown. Leg I dark (Araneae: Nemesiidae) burrow (Forster 1995). Periegops keani does not appear to build a web for prey capture and 18 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae) probably builds silken retreats, like P. suterii. The lack be located at East Cape, 28S sequence data could confirm of webs suggests that it is a cursorial, night-time hunter. whether they were the same species. Adults have been found in March and September, but Forster & Forster (1999) stated that they had ex- given that the climates it is found in are milder than those amined a female specimen from the Aldermen Islands. of P. suterii, it seems likely that adults occur throughout This specimen was collected from Ruamahuaiti Island in the year. November 1972 and loaned to Ray Forster from NZAC DNA. Three cytochrome c oxidase subunit 1 and two 28S (G. Hall, pers. comm.). We have examined this specimen rRNA sequences for this species are listed in GenBank and determined it to be a subadult male, as it has swollen (www.ncbi.nlm.nih.gov/GenBank/) under accession palpal tibiae, which are characteristic of males in their numbers JX174288–JX174290, JX174292, and JX174293. final moult before adulthood. The specimen did not have an epigyne or internal genitalia and we believe Forster Etymology. The specific name is in honour of the senior & Forster (1999) made a typographical error in stating author’s friend and colleague, John Kean, who helped the specimen was female. We consider that this speci- collect the type specimen. men is P. keani. Ruamahuaiti Island is only 2 km from Remarks. The bulb of the holotype was not oriented to Ruamahuanui Island and the land below the two islands the same angle with respect to the cymbium (Fig. 9) as is no more than 20 m under the sea (Hayward & Moore the specimens illustrated of P. suterii (Fig. 6, 7) and P. 1973), therefore, the two populations would have been australia (Fig. 8); however, when oriented in a slight connected up until 9000 years ago (Schofield & Thomp- dorsal direction (Fig. 9a), it looks the same as those of P. son 1964; McKinnon 1997). suterii, other than the flattened flanges near the embolus Size may help differentiate P. keani from P. sute- tip, which are not as pronounced in P. keani. rii even though we have only examined a single adult We consider that a female specimen found in for- specimen of the former species. The subadult male from est below the East Cape Lighthouse is P. keani. Forster Ruamahuaiti Island is larger (body length = 8.8 mm, (1995) remarked that the genitalia of this specimen was carapace length = 4.3 mm) than any of the 11 adult male indistinguishable from that of P. suterii. The female P. suterii that we have examined. Also, a P. keani speci- specimen from East Cape was collected by Grace Hall men (MONZ AS.002301) that appeared to be one or two (NZAC) on 30 September 1993 and was loaned to the moults from adulthood was larger (body length = 9.3, late Ray Forster, but has not been returned to NZAC (G. carapace length = 3.9) than five of the 11 adult specimens Hall pers. comm.) and it has not been located at OMNZ of P. suterii examined; based on this and the size of the (C. Fraser, pers. comm.). Unsuccessful searches for fur- male, it would be reasonable to expect an adult female P. ther specimens have been conducted in the small (~10 keani to be larger than any P. suterii female. hectare) patch of forest below the East Cape Lighthouse in October 1994 (G. Hall), February 1995 (G. Hall), Sep- tember 1995 (G. Hall & P. J. Sirvid), November 1995 (G. Hall & L. J. Boutin), December 2009 (G. Hall), Novem- ber 2010 (J. Malumbres-Olarte) and February 2012 (C. J. Vink). Searches of the forest floor (including rock and log turning) were conducted in all trips, Stanwellia sp. burrows were searched in most trips, and pitfall trapping and litter searches were also conducted in the former and later trips. Periegops keani may conceivably now be ex- tinct from that site. It is possible that the population at East Cape could be a separate species to P. keani, as Periegops probably has limited dispersal ability. Although the Aldermen Is- lands and East Cape are 233 km apart, the Aldermen Is- lands would have been connected to the mainland from the Last Glacial Maximum (McKinnon 1997; Trewick & Bland 2012) up until 12000 years ago (Schofield & Thompson 1964; McKinnon 1997) and there would have been continuous forest between the two sites (McKinnon 1997; Trewick & Bland 2012). If a fresh specimen could Fauna of New Zealand 70 ­19­

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APPENDIX A: Glossary of technical terms maxilla — (= endite) the expanded lobe of the palpal coxa situated laterally of the labium abdomen — posterior division of the spider body; (plural maxillae) sometimes referred to as the opisthosoma metatarsus — sixth segment of the legs; absent in bulb — refers to the male pedipalpal organ as a the pedipalps whole paturon — the basal segment of a chelicerae carapace — the hard dorsal covering of the pedipalp — six-segmented second appendage of cephalothorax the cephalothorax, anterior to legs I cephalothorax — anterior division of the spider patella — fourth segment of the legs and pedipalp body; sometimes referred to as the prosoma promarginal — anterior margin chelicerae — first pair of appendages of the proximal — near the base cephalothorax, consisting of two segments retrolateral — on the outer side i.e., the surface (the distal segment is called the fang, basal nearer to the posterior end of the body segment is called the paturon) retromarginal — posterior margin coxa — first or basal segment of the legs sclerotised — hardened by sclerotin or other cymbium — tarsus of the adult male pedipalp substances in the cuticle distal — near the apex seta — a sclerotised hair-like projection arising dorsal — upper (surface) from the cuticle (plural setae) epigastric furrow — a transverse groove across sternum — plate on the ventral surface of the the anterior ventral part of the abdomen cephalothorax between the coxae of the legs epigyne — the sclerotised region of female spiders covering the internal genitalia tarsus — last segment of the legs and pedipalp and located between the book lungs and tibia — fifth segment of legs and pedipalp anterior of the epigastric furrow trochanter — second segment of the leg and embolus — the intromittent structure of the bulb pedipalp containing the terminal portion of the ventral — lower (surface) ejaculatory duct femur — third segment of the legs and pedipalps fovea — depression on the thoracic region of the carapace where muscles of sucking stomach are attached internally haplogyne — the primitive form of spider genitalia where the female has the copulatory openings internally, within the gonopore, and typically lacks a sclerotised epigyne, and the male has relatively simple pedipalps. This form is found in the basal spiders, including Mygalomorphae, Gradungulidae and Haplogynae. 24 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

APPENDIX B: Collection details of specimens serve, 43°48.59’S, 173°01.28’E, 11 Oct 2002, C. J. examined. Localities (including coordinates) and Vink, AMNH ARAMR-000616; 1 ♀, Hinewai Reserve, dates collected, collectors, and institutions of 43°48.6’S, 173°01.3’E, 27 Aug 1996, C. J. Vink, specimens examined. LUNZ 00012734; 1 ♂, Hinewai Reserve, 43°48.69’S, 173°00.95’E, 11 Jan 2011, M. H. Bowie, LUNZ Periegops suterii 00012735; 1 ♀, Akaroa, 43°48.8’S, 172°57.5’E, 16 Oct 1920, G. Archey, OMNZ IV35928; 1 subadult ♀, MC. 3 ♂, 1 ♀, Riccarton Bush, 43°31.7’S, 172°35.7’E, Fishermans Bay , 43°49.54’S, 173°03.85’E, 21 Dec 9 Apr 1994, A. D. Blest, OMNZ IV35925; 1 ♂, Ken- 2010, M. H. Bowie, LUNZ 00012736. nedys Bush Reserve, 43°37.9’S, 172°37.3’E, 22 Apr 2011, C. J. Vink, H. P. & L. J. Hudson Vink, LUNZ Periegops keani 00012715; 1 ♀, Kennedys Bush Reserve, 43°37.9’S, 172°37.3’E, 13 Aug 2011, C. J. Vink & H. P. Hudson CL. 1 immature, Ruamahuanui Island, Vink, LUNZ 00012718; 1 ♀, Kennedys Bush Reserve, 36°57.3'S,176°05.5'E, 7 Nov 2002, M. D. Wake- 43°37.9’S, 172°37.3’E, 13 Aug 2011, C. J. Vink & H. lin, MONZ AS.002302; 1 immature, Ruamahuanui P. Hudson Vink, LUNZ 00012719; 1 ♂, Rhodes Bush, Island, 36°57.25'S, 176°05.52'E, 19 Nov 2003, 43°40’S, 172°37’E, 13 Nov 1915, G. Archey, OMNZ B. M. Fitzgerald, MONZ AS.002310; 1 immature, IV35927; 1 ♀, Hay Reserve, 43°42.18’S, 172°53.87’E, Ruamahuanui Island, 36°57.25'S, 176°05.52'E, 13 Mar 2003, C. J. Vink, LUNZ 00012720; 2 ♀, Kai- 19 Nov 2003, B. M. Fitzgerald, MONZ AS.002300; tuna Valley, 43°43’S, 172°45’E, 11 Sep 1949, R. R. holotype ♂, Ruamahuanui Island, 36°57.25'S, Forster, OMNZ IV35923; 1 ♂, Montgomery Park 176°05.52'E, 23 Mar 2012, C. J. Vink & J. M. Kean, Reserve, 43°44.7’S, 172°52.2’E, 5 Nov 2010, C. J. LUNZ 00012716; 1 immature, Ruamahuanui Island, Vink & J. Malumbres-Olarte, LUNZ 00012721; 1 ♂, 36°57.25'S, 176°05.52'E, 23 Mar 2012, C. J. Vink & J. Montgomery Park Reserve, 43°44.7’S, 172°52.2’E, M. Kean, LUNZ 00012717; 1 subadult ♀, 1 immature, 5 Nov 2010, C. J. Vink & J. Malumbres-Olarte, Ruamahuanui Island, 36°57.25'S, 176°05.52'E, 18 LUNZ 00012722; 1 ♂, Montgomery Park Reserve, Apr 2005, B. M. Fitzgerald & P. MacDonald, MONZ 43°44.7’S, 172°52.2’E, 5 Nov 2010, C. J. Vink & J. AS.002301; 1 subadult ♂, Ruamahuaiti Island, Malumbres-Olarte, LUNZ 00012723; 1 ♂, Mont- 36°58.5'S, 176°05'E, 8-12 November 1972, G. W. gomery Park Reserve, 43°44.7’S, 172°52.2’E, 5 Ramsay, NZAC. Nov 2010, J. Malumbres-Olarte & C. J. Vink, LUNZ 00012724; 1 ♀, 1 subadult ♀, Montgomery Park Reserve, 43°44.7’S, 172°52.2’E, 5 Nov 2010, J. Malumbres-Olarte & C. J. Vink, LUNZ 00012725; 1 ♀, Panama Rock , 43°44.77’S, 173°02.49’E, 12 Jan 2011, M. H. Bowie, LUNZ 00012726; 1 ♀, Panama Rock , 43°44.77’S, 173°02.49’E, 12 Jan 2011, M. H. Bowie, LUNZ 00012727; 1 ♂, Otepa- totu Reserve, 43°44.90’S, 173°00.95’E, 4 Jan 2011, M. H. Bowie, LUNZ 00012728; 1 ♂, Otepatotu Reserve, 43°44.90’S, 173°00.95’E, 12 Jan 2011, M. H. Bowie, LUNZ 00012729; 1 ♀, Otepatotu Re- serve, 43°44.90’S, 173°00.95’E, 4 Jan 2011, M. H. Bowie, LUNZ 00012730; 1 ♀, Otepatotu Reserve, 43°44.93’S, 173°00.95’E, 12 Jan 2011, M. H. Bowie, LUNZ 00012731; 1 ♂, Tititipounamu , 43°45.47’S, 173°01.44’E, 4 Jan 2011, M. H. Bowie, LUNZ 00012732; 1 ♀, Little River, 43°46’S, 172°47’E, 10 Jan 1985, A. C. Harris, OMNZ IV35924; 1 ♂, Ellan- gowan Reserve, 43°47.89’S, 173°02.08’E, 12 Jan 2011, M. H. Bowie, LUNZ 00012733; 1 ♂, Hinewai Reserve, 43°48.59’S, 173°01.28’E, 11 Oct 2002, C. J. Vink, AMNH ARAMR-000615; 1 ♀, Hinewai Re- Fauna of New Zealand 70 ­25­

ILLUSTRATIONS

Fig. 1 Schematic dorsal view, Periegops suterii, female (LUNZ 00012720). Actual size on the right. 26 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Fig. 2, 3 Periegops keani (holotype): (2) anterior view of chelicerae; (3) ventral view of mouthparts. Fauna of New Zealand 70 ­27­

Fig. 4, 5 Retrolateral view of leg I: (4) Periegops suterii (LUNZ 00012722); (5) Periegops keani (holotype). 28 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Fig. 6–9 Retrolateral view of left male pedipalps (b – bulb; c – cymbium; e – embolus), arrow indicates dorsal projection on the cymbium: (6) Periegops suterii, Kennedys Bush Reserve (LUNZ 00012715); (7) Periegops suterii, Montgomery Park Reserve (LUNZ 00012721); (8) Periegops australia, Mt Goonaneman (QMB S20421); (9) Periegops keani, Ruamahuanui Island (holotype) (9a = laterodorsal view of bulb and cymbium). Fauna of New Zealand 70 ­29­

Fig. 10, 11 Ventral view of female epigastric area: (10) Periegops suterii, Hay Reserve (LUNZ 00012720); (11) Periegops australia, Mt Goonaneman (QMB S20419). 30 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Fig. 12, 13 Dorsal view of female internal genitalia (p – poreplate; ic – invaginated cup): (12) Periegops suterii, Hay Reserve (LUNZ 00012720); (13) Periegops australia, Mt Goonaneman (QMB S20419). Fauna of New Zealand 70 ­31­

Fig. 14 Dorsal view of Periegops suterii, male (LUNZ 00012721). Actual size on the right. 32 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Fig. 15 Dorsal view of Periegops keani, male (holotype). Actual size on the right. Fauna of New Zealand 70 ­33­

Fig. 16 Photograph of Periegops suterii, female (LUNZ 00012718). Photographer: Bryce McQuillan. 34 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

DISTRIBUTION MAPS

Map 1 Collection localities, Periegops suterii. Inset shows localities on and near Banks Peninsula, and labelled locations indicate collection sites for DNA specimens.

Map 2 Collection localities, Periegops keani.

Map 2 Collection localities, Periegops keani. Fauna of New Zealand 70 ­35­

TAXONOMIC INDEX

Page numbers in bold type denote a description, and Periegopidae this contribution in italic type illustrations. A suffixed letter ‘k’ indicates Periegops this contribution a key, and ‘m’ a map. Periegops australia 8, 10, 15, 17, 18, 28, 29, 30 Periegops hirsutus 8, 15, 16, 17 australia, Periegops 8, 10, 15, 17, 18, 28, 29, 30 Periegops keani 7, 11, 12, 13, 15k, 17–18, 24, 26, 27, 28, 32, 34m Caponiidae 9 Periegops suteri 15, 17 Cephalobidae 11 Periegops suterii 5, 7, 8, 9, 10, 11, 12, 13, 14, 15k, 15–17, 18, 24, 25, 27, 28, 29, 30, 31, 33, 34m Diguetidae 9 Pholcidae 9 Drymusa 9 Plectreuridae 9 Drymusidae 9 Dysderidae 8 Segestria 8 Segestria suterii 15, 16, 17 Entelegynae 14 Scytodidae 9 Scytodoidea 9 Haplogynae 9 Sicariidae 9 Hemideina 14 Sicarioidea 9 hirsutus, Periegops 8, 15, 16, 17 Stanwellia sp. 17, 18 Hypochilidae 14 suteri, Periegops 15, 17 suterii, Periegops 5, 7, 8, 9, 10, 11, 12, 13, 14, 15k, keani, Periegops 7, 11, 12, 13, 15k, 17–18, 24, 26, 15–17, 18, 24, 25, 27, 28, 29, 30, 31, 33, 34m 27, 28, 32, 34m suterii, Segestria 15, 16, 17 Leptonetidae 9 Telemidae 9 Nemesiidae 17 Tetrablemmidae 9

Ochyroceratidae 9 36 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae) Fauna of New Zealand 70 ­37­ 38 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Fauna of New Zealand publications

1 Terebrantia (Insecta: Thysanoptera). Laurence A. Mound 21 Margarodidae (Insecta: Hemiptera). C. F. Morales. ISBN & Annette K. Walker. ISBN 0-477-06687-9, 23 December 0-477-02607-9, 27 May 1991, 124 pp...... $34.95 1982, 120 pp...... $29.95 22 Notonemouridae (Insecta: Plecoptera). I. D. McLellan, 2 Osoriinae (Insecta: Coleoptera: Staphylinidae). H. Pauline ISBN 0-477-02518-8, 27 May 1991, 64 pp...... $24.95 McColl. ISBN 0-477-06688-7, 23 December 1982, 96 23 Sciapodinae, Medeterinae (Insecta: Diptera) with a pp...... $18.60 generic review of the Dolichopodidae. D. J. Bickel. ISBN 3 Anthribidae (Insecta: Coleoptera). B. A. Holloway. 0-477-02627-3, 13 January 1992, 74 pp...... $27.95 ISBN 0-477-06703-4, 23 December 1982, 272 pp...... 24 Therevidae (Insecta: Diptera). L. Lyneborg. ISBN 0-477- ...... $41.00 02632-X, 4 March 1992, 140 pp...... $34.95 4 Eriophyoidea except Eriophyinae (Arachnida: Acari). D. 25 Cercopidae (Insecta: Homoptera). K. G. A. Hamilton C. M. Manson. ISBN 0-477-06745-X,12 November 1984, & C. F. Morales. ISBN 0-477-02636-2, 25 May 1992, 40 144 pp...... $29.95 pp...... $17.95 5 Eriophyinae (Arachnida: Acari: Eriophyoidea). D. C. M. 26 Tenebrionidae (Insecta: Coleoptera): catalogue of types Manson. ISBN 0-477-06746-8, 14 November 1984, 128 and keys to taxa. J. C. Watt. ISBN 0-477-02639-7, 13 July pp...... $29.95 1992, 70 pp...... $27.95 6 Hydraenidae (Insecta: Coleoptera). R. G. Ordish. ISBN 27 Antarctoperlinae (Insecta: Plecoptera). I. D. McLellan. 0-477-06747-6, 12 November 1984, 64 pp...... $18.60 ISBN 0-477-01644-8, 18 February 1993, 70 pp. . $27.95 7 Cryptostigmata (Arachnida: Acari) — a concise review. 28 Larvae of Curculionoidea (Insecta: Coleoptera): a M. Luxton. ISBN 0-477-06762-X, 8 December 1985, 112 systematic overview. Brenda M. May. ISBN 0-478-04505-0, pp...... $29.95 14 June 1993, 226 pp...... $55.00 8 Calliphoridae (Insecta: Diptera). James P. Dear. ISBN 29 Cryptorhynchinae (Insecta: Coleoptera: Curculionidae). 0-477-06764-6. 24 February 1986, 88 pp...... $18.60 C. H. C. Lyal. ISBN 0-478-04518-2, 2 December 1993, 308 (Insecta). S. L. Tuxen. ISBN 0-477-06765-4, 24 9 Protura pp...... $65.00 February 1986, 52 pp...... $18.60 30 Hepialidae (Insecta: Lepidoptera). J. S. Dugdale. ISBN (Insecta: Thysanoptera). Laurence A. Mound 10 Tubulifera 0-478-04524-7, 1 March 1994, 164 pp...... $42.50 & Annette K. Walker. ISBN 0-477-06784-0, 22 September 1986, 144 pp...... $34.65 31 Talitridae (Crustacea: Amphipoda). K. W. Duncan. ISBN 0-478-04533-6, 7 October 1994, 128 pp...... $36.00 11 Pseudococcidae (Insecta: Hemiptera). J. M. Cox. ISBN 0-477-06791-3, 7 April 1987, 232 pp...... $49.95 32 Sphecidae (Insecta: Hymenoptera). A. C. Harris, ISBN 0-478-04534-4, 7 October 1994, 112 pp...... $33.50 12 Pompilidae (Insecta: Hymenoptera). A. C. Harris. ISBN 0-477-02501-3, 13 November 1987, 160 pp...... $39.95 33 Moranilini (Insecta: Hymenoptera). J. A. Berry. ISBN 0-478-04538-7, 8 May 1995, 82 pp...... $29.95 13 Encyrtidae (Insecta: Hymenoptera). J. S. Noyes. ISBN 0-477-02517-X, 9 May 1988, 192 pp...... $44.95 34 Anthicidae (Insecta: Coleoptera). F. G. Werner & D. S. Chandler. ISBN 0-478-04547-6, 21 June 1995, 64 pp. .. 14 Lepidoptera — annotated catalogue, and keys to family-group taxa. J. S. Dugdale. ISBN 0-477-02518-8, ...... $26.50 23 September 1988, 264 pp...... $49.95 35 Cydnidae, Acanthosomatidae, and Pentatomidae 15 Ambositrinae (Insecta: Hymenoptera: Diapriidae). I. D. (Insecta: Heteroptera): systematics, geographical Naumann. ISBN 0-477-02535-8, 30 December 1988, 168 distribution, and bioecology. M.-C. Larivière. ISBN 0-478- pp...... $39.95 09301-2, 23 November 1995, 112 pp...... $42.50 16 Nepticulidae (Insecta: Lepidoptera). Hans Donner & 36 Leptophlebiidae (Insecta: Ephemeroptera). D. R. Towns Christopher Wilkinson. ISBN 0-477-02538-2, 28 April 1989, & W. L. Peters. ISBN 0-478-09303-9, 19 August 1996, 144 92 pp...... $22.95 pp...... $39.50 17 Mymaridae (Insecta: Hymenoptera) — introduction, and 37 Coleoptera: family-group review and keys to review of genera. J. S. Noyes & E. W. Valentine. ISBN identification. J. Klimaszewski & J. C. Watt. ISBN 0-478- 0-477-02542-0, 28 April 1989, 100 pp...... $24.95 09312-8, 13 August 1997, 199 pp...... $49.50 18 Chalcidoidea (Insecta: Hymenoptera) — introduction, 38 Naturalised terrestrial Stylommatophora (Mollusca: and review of genera in smaller families. J. S. Noyes & Gastropoda). G. M. Barker. ISBN 0-478-09322-5, 25 E. W. Valentine. ISBN 0-477-02545-5, 2 August 1989, 96 January 1999, 253 pp...... $72.50 pp...... $24.95 39 Molytini (Insecta: Coleoptera: Curculionidae: Molytinae). 19 Mantodea (Insecta), with a review of aspects of functional R. C. Craw. ISBN 0-478-09325-X, 4 February1999, 68 morphology and biology. G. W. Ramsay. ISBN 0-477- pp...... $29.50 02581-1, 13 June 1990, 96 pp...... $24.95 40 Cixiidae (Insecta: Hemiptera: Auchenorrhyncha). M.-C. 20 Bibionidae (Insecta: Diptera). Roy A. Harrison. ISBN Larivière. ISBN 0-478-09334-9, 12 November 1999, 93 0-477-02595-1. 13 November 1990, 28 pp...... $14.95 pp...... $37.50 Fauna of New Zealand 70 ­39­

41 Coccidae (Insecta: Hemiptera: Coccoidea). C. J. 60 Carabidae (Insecta: Coleoptera): synopsis of Hodgson & R. C. Henderson. ISBN 0-478-09335-7, 23 supraspecific taxa. A. Larochelle & M.-C. Larivière. ISBN February 2000, 264 pp...... $72.50 978-0-478-09394-0, 21 November 2007, 188 pp. . $54.00 42 Aphodiinae (Insecta: Coleoptera: Scarabaeidae). Z. T. 61 Lucanidae (Insecta: Coleoptera). B. A. Holloway. ISBN Stebnicka. ISBN 0-478-09341-1, 15 June 2001, 64 pp. . 978-0-478-09395-7, 21 November 2007, 254 pp. . $75.00 ...... $29.50 62 Trechini (Insecta: Coleoptera: Carabidae: Trechinae). 43 Carabidae (Insecta: Coleoptera): catalogue. A. J. I. Townsend. ISBN 978-0-478-34717-9 (print), 978-0- Larochelle & M.-C. Larivière. ISBN 0-478-09342-X, 15 478-34716-6 (online), 16 June 2010, 101 pp. .... $49.50 June 2001, 285 pp...... $72.50 63 Auchenorrhyncha (Insecta: Hemiptera): catalogue. 44 Lycosidae (Arachnida: Araneae). C. J. Vink. ISBN 0-478- M.-C. Larivière, M. J. Fletcher & A. Larochelle. ISBN 09347-0, 23 December 2002, 94 pp...... $37.50 978-0-478-34720-3 (print), 978-0-478-34721-0 (online), 45 Nemonychidae, Belidae, Brentidae (Insecta: 16 June 2010, 232 pp...... $75.00 Coleoptera: Curculionoidea). G. Kuschel. ISBN 0-478- 64 Pisauridae (Arachnida: Araneae). C. J. Vink & N. 09348-9, 28 April 2003, 100 pp...... $40.00 Dupérré. ISBN 978-0-478-34722-7 (print), 978-0-478- 46 Nesameletidae (Insecta: Ephemeroptera). Terry R. 34723-4 (online), 13 July 2010, 60 pp...... $37.50 Hitchings & Arnold H. Staniczek. ISBN 0-478-09349-7, 65 Izatha (Insecta: Lepidoptera: Gelechioidea: Oeco­ 14 May 2003, 72 pp...... $32.50 phoridae). Robert J. B. Hoare. ISBN 978-0-478-34724-1 47 Erotylidae (Insecta: Coleoptera: Cucujoidea): phylogeny (print), 978-0-478-34725-8 (online), 2 September 2010, and review. R. A. B. Leschen. ISBN 0-478-09350-0, 5 June 201 pp...... $75.00 2003,108 pp...... $42.50 66 Diaspididae (Insecta: Hemiptera: Coccoidea). R. C. 48 Scaphidiinae (Insecta: Coleoptera: Staphylinidae). I. Henderson. ISBN 978-0-478-34726-5 (print), 978-0-478- Löbl & R. A. B. Leschen. ISBN 0-478-09353-5,18 November 34727-2 (online), 23 May 2011, 275 pp...... $89.00 2003, 94 pp...... $37.50 67 Peloridiidae (Insecta: Hemiptera: Coleorrhyncha). M.-C. 49 Lithinini (Insecta: Lepidoptera: Geometridae: Larivière, D. Burckhardt & A. Larochelle. ISBN 978-0-478- Ennominae). J. D. Weintraub & M. J. Scoble. ISBN 0-478- 34730-2 (print), 978-0-478-34731-9 (online), 14 November 09357-8, 29 April 2004, 48 pp...... $24.50 2011, 78 pp...... $48.00 50 Heteroptera (Insecta: Hemiptera): catalogue. M.-C. 68 Simuliidae (Insecta: Diptera). Douglas A. Craig, Ruth Larivière & A. Larochelle. ISBN 0-478-09358-6, 14 May E. G. Craig & Trevor K. Crosby. ISBN 978-0-478-34734- 2004, 330 pp...... $89.00 0 (print), 978-0-478-34735-7 (online), 29 June 2012, 336 51 Coccidae (Insecta: Hemiptera: Coccoidea): adult males, pp...... $95.00 pupae and prepupae of indigenous species. C. J. Hodgson 69 Carabidae (Insecta: Coleoptera): synopsis of species, & R. C. Henderson. ISBN 0-478-09360-8, 22 June 2004, Cicindelinae to Trechinae (in part) . A. Larochelle & M.-C. 228 pp...... $65.00 Larivière. ISBN 978-0-478-34738-8 (print), 978-0-478- 52 Raphignathoidea (Acari: Prostigmata). Qing-Hai Fan 34739-5 (online), 7 March 2013, 193 pp...... $75.00 & Zhi-Qiang Zhang. ISBN 0-478-09371-3, 20 May 2005, 70 Periegopidae (Arachnida: Araneae). C. J. Vink, N. 400 pp...... $89.00 Dupérré & J. Malumbres-Olarte. ISBN 978-0-478-34740- 53 Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae). 1 (print), 978-0-478-34741-8 (online), 7 March 2013, 41 A. Larochelle & M.-C. Larivière. ISBN 0-478-09369-1, 4 pp...... $29.00 July 2005, 160 pp...... $55.00 54 Hierodoris (Insecta: Lepidoptera: Gelechoidea: Oeco­ Visit the Manaaki Whenua Press Website at: phoridae), and overview of Oecophoridae. Robert J. B. http://www.mwpress.co.nz/ Hoare. ISBN 0-478-09378-0, 24 December 2005, 100 for further information. pp...... $40.00 To access on-line extracts and PDFs from this series 55 Criconematina (Nematoda: Tylenchida). W. M. Wouts. ISBN 0-478-09381-0, 24 March 2006, 232 pp. ... $65.00 visit: http://fnz.landcareresearch.co.nz/ 56 Tyrophagus (Acari: Astigmata: Acaridae). Qing-Hai Fan & Zhi-Qiang Zhang. ISBN 978-0-478-09386-5, 4 June 2007, 291 pp...... $80.00 57 Apoidea (Insecta: Hymenoptera). B. J. Donovan. ISBN 978-0-478-09389-6, 7 September 2007, 295 pp .. $89.00 58 Alysiinae (Insecta: Hymenoptera: Braconidae). J. A. Berry. ISBN 978-0-478-09390-2, 7 September 2007, 95 pp...... $45.00 59 Erotylinae (Insecta: Coleoptera: Cucujoidea: Erotylidae): taxonomy and biogeography. Paul E. Skelley & Richard A. B. Leschen. ISBN 978-0-478-09391-9, 7 September 2007, 59 pp...... $30.00 40 Vink, Dupérré & Malumbres-Olarte (2013): Periegopidae (Arachnida: Araneae)

Taxonomic groups covered in the Pseudococcidae (J. M. Cox, FNZ 11, 1987) Fauna of New Zealand series Peloridiidae (M.-C. Larivière, D. Burckhardt & A. Larochelle, FNZ 67, 2011). Insecta Coleoptera Hymenoptera Family-group review and keys to identification (J. Klimaszew- Apoidea (B. J. Donovan, FNZ 57, 2007) ski & J. C. Watt, FNZ 37, 1997) Braconidae: Alysiinae (J. A. Berry, FNZ 58, 2007) Anthribidae (B. A. Holloway, FNZ 3, 1982) Chalcidoidea: introduction, and review of smaller families (J. Anthicidae (F. G. Werner & D. S. Chandler, FNZ 34, 1995) S. Noyes & E. W. Valentine, FNZ 18, 1989) Carabidae: catalogue (A. Larochelle & M.-C. Larivière, FNZ Diapriidae: Ambositrinae (I. D. Naumann, FNZ 15, 1988) 43, 2001); synopsis of supraspecific taxa (A. Larochelle Encyrtidae (J. S. Noyes, FNZ 13, 1988) & M.-C. Larivière, FNZ 60, 2007); synopsis of species, Mymaridae (J. S. Noyes & E. W. Valentine, FNZ 17, 1989) Cicindelinae to Trechinae (in part) (A. Larochelle & M.-C. Pompilidae (A. C. Harris, FNZ 12, 1987) Larivière, FNZ 69, 2013) Pteromalidae: Eunotinae: Moranilini (J. A. Berry, FNZ 33, Carabidae: Harpalinae: Harpalini (A. Larochelle & M.-C. 1995) Larivière, FNZ 53, 2005) Sphecidae (A. C. Harris, FNZ 32, 1994) Carabidae: Trechinae: Trechini (J. I. Townsend, FNZ 62, Lepidoptera 2010) Annotated catalogue, and keys to family-group taxa (J. S. Curculionidae: Cryptorhynchinae (C. H. C. Lyal, FNZ 29, Dugdale, FNZ 14, 1988) 1993) Geometridae: Ennominae: Lithinini (Jason D. Weintraub & Curculionidae: Molytinae: Molytini (R. C. Craw, FNZ 39, Malcolm J. Scoble, FNZ 49, 2004) 1999) Hepialidae (J. S. Dugdale, FNZ 30, 1994) Curculionoidea: Nemonychidae, Belidae, Brentidae (G. Ku- Nepticulidae (Hans Donner & Christopher Wilkinson, FNZ schel, FNZ 45, 2003) 16, 1989) Curculionoidea larvae: a systematic overview (Brenda M. Oecophoridae: Hierodoris (Robert J. B. Hoare, FNZ 54, May, FNZ 28, 1993) 2005); Izatha (Robert J. B. Hoare, FNZ 65, 2010). Erotylidae: phylogeny and review (Richard A. B. Leschen, FNZ 47, 2003); Erotylinae: taxonomy and biogeography Mantodea, with a review of aspects of functional morphology (Paul E. Skelley & Richard A. B. Leschen, FNZ 59, 2007) and biology (G. W. Ramsay, FNZ 19, 1990) Hydraenidae (R. G. Ordish, FNZ 6, 1984) Lucanidae (B. A. Holloway, FNZ 61, 2007) Plecoptera Antarctoperlinae (I. D. McLellan, FNZ 27, 1993) Scarabaeidae: Aphodiinae (Z. T. Stebnicka, FNZ 42, 2001) Notonemouridae (I. D. McLellan, FNZ 22, 1991) Staphylinidae: Osoriinae (H. Pauline McColl, FNZ 2, 1982) Staphylinidae: Scaphidiinae (I. Löbl & Richard A. B. Leschen, Protura (S. L. Tuxen, FNZ 9, 1986) FNZ 48, 2003) Tenebrionidae: catalogue of types and keys to taxa (J. C. Thysanoptera Watt, FNZ 26, 1992) Terebrantia (Laurence A. Mound & Annette K. Walker, FNZ 1, 1982) Diptera Tubulifera (Laurence A. Mound & Annette K. Walker, FNZ Bibionidae (Roy A. Harrison, FNZ 20, 1990) 10, 1986) Calliphoridae (James P. Dear, FNZ 8, 1986) Arachnida Dolichopodidae: Sciapodinae, Medeterinae with a generic Acari review (D. J. Bickel, FNZ 23, 1992) Acaridae: Tyrophagus (Qing-Hai Fan & Zhi-Qiang Zhang, Simuliidae (Douglas A. Craig, Ruth E. G. Craig, Trevor K. FNZ 56, 2007) Crosby, FNZ 68, 2012) Cryptostigmata — a concise review (M. Luxton, FNZ 7, 1985) Therevidae (L. Lyneborg, FNZ 24, 1992) Eriophyoidea except Eriophyinae (D. C. M. Manson, FNZ 4, Ephemeroptera 1984) Leptophlebiidae (D. R. Towns & W. L. Peters, FNZ 36, 1996) Eriophyinae (D. C. M. Manson, FNZ 5, 1984) Nesameletidae (Terry R. Hitchings & Arnold H. Staniczek, Raphignathoidea (Qing-Hai Fan & Zhi-Qiang Zhang, FNZ FNZ 46, 2003) 52, 2005) Araneae Hemiptera Lycosidae (C. J. Vink, FNZ 44, 2002) Auchenorrhyncha: catalogue (M.-C. Larivière, M. J. Fletcher Periegopidae (C. J. Vink, N. Dupérré & Malumbres-Olarte, & A. Larochelle, FNZ 63, 2010) FNZ 70, 2013 Cercopidae (K. G. A. Hamilton & C. F. Morales, FNZ 25, Pisauridae (C. J. Vink & N. Dupérré, FNZ 64, 2010) 1992) Cixiidae (M.-C. Larivière, FNZ 40, 1999) Crustacea Coccidae (C. J. Hodgson & R. C. Henderson, FNZ 41, 2000); Amphipoda adult males, pupae and prepupae of indigenous species Talitridae (K. W. Duncan, FNZ 31, 1994) (C. J. Hodgson & R. C. Henderson, FNZ 51, 2004) Cydnidae, Acanthosomatidae, and Pentatomidae (M.-C. Mollusca Larivière, FNZ 35, 1995) Gastropoda Diaspididae (R. C. Henderson, FNZ 66, 2011) Naturalised terrestrial Stylommatophora (G. M. Barker, FNZ Heteroptera: catalogue (M.-C. Larivière & A. Larochelle, FNZ 38, 1999) 50, 2004) Nematoda Margarodidae (C. F. Morales, FNZ 21, 1991) Tylenchida: Criconematina (W. M. Wouts, FNZ 55, 2006) Fauna of New Zealand 70 ­41­

NOTICES NGĀ PĀNUI

This series of refereed publications has been established to Kua whakatūria tēnei huinga pukapuka hei whakahauhau i encourage those with expert knowledge to publish concise ngā tohunga whai mātauranga kia whakaputa i ngā kōrero yet comprehensive accounts of elements in the New Zea- poto, engari he whaikiko tonu, e pā ana ki ngā aitanga land fauna. The series is professional in its conception and pepeke o Aotearoa. He tōtika tonu te āhua o ngā tuhituhi, presentation, yet every effort is made to provide res­ources engari ko te tino whāinga, kia mārama te marea ki ngā for identification and information that are accessible to tohu tautuhi o ia ngārara, o ia ngārara, me te roanga atu o the non-specialist. ngā kōrero mō tēnā, mō tēnā.

Fauna of N.Z. deals with non-marine invertebrates only, He titiro whāiti tā tēnei pukapuka ki ngā mea noho whenua, since the vertebrates are well documented, and marine kāore he tuarā; i pēnei ai i te mea kei te mōhio whānuitia forms are covered by the series NIWA Biodiversity ngā mea whai tuarā, ā, ko ngā mea noho moana, koirā te Memoirs. tino kaupapa o te huinga pukapuka NIWA Biodiversity Memoirs. Contributions are invited from any person with the requisite specialist skills and resources. Material from the Ka āhei te tangata ki te whakauru tuhituhinga mehemea N.Z. Arthropod Collection is available for study. kei a ia ngā tohungatanga me ngā rauemi e tutuki pai ai tana mahi. Heoi anō, e wātea ana te Kohinga Angawaho Contributors should discuss their intentions with a member o Aotearoa hei āta tirotiro mā te tangata mehemea he of the Editorial Board or with the Series Editor before āwhina kei reira. commencing work; all necessary guidance will be given. Me whāki te kaituhi i ōna whakaaro ki tētahi o te Kāhui Subscribers should address enquiries to Fauna of N.Z., Ārahi Whakarōpūtanga Tuarā-Kore, ki te Ētita rānei i Manaaki Whenua Press, Landcare Research, P.O. Box 40, mua i te tīmatanga, ā, mā rātou a ia e ārahi mō te wāhi ki Lincoln 7640, New Zealand. tana tuhinga.

Ko te hunga pīrangi hoko pukapuka, me tuhi ki Fauna of N.Z., Manaaki Whenua Press, Manaaki Whenua, Pouaka Poutāpeta 40, Lincoln 7640, Aotearoa.