Zootaxa 3637 (5): 521–540 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3637.5.2 http://zoobank.org/urn:lsid:zoobank.org:pub:447D8DF5-F922-4B3A-AC43-A85225E56C57 New species of Mouse (Araneae: : : ) from the Pilbara region, Western Australia

DANILO HARMS1, 2, 3, 4 & VOLKER W. FRAMENAU1, 2, 3 1 School of Biology, The University of Western Australia, 35 Stirling Highway, Crawley, Western Australia 6009, Australia. 2 Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia. 3 Phoenix Environmental Sciences Pty Ltd, 1/511 Wanneroo Road, Balcatta, Western Australia 6021, Australia. 4 Corresponding author. E-mail: [email protected]

Abstract

Two new species of Mouse Spiders, Missulena, from the Pilbara region in Western Australia are described based on morphological features of males. Missulena faulderi sp. nov. and Missulena langlandsi sp. nov. are currently known from a small area in the southern Pilbara only. Mitochondrial cytochrome c oxidase subunit I (COI) sequence divergence failed in clearly delimiting species in Missulena, but provided a useful, independent line of evidence for taxonomic work in addition to morphology.

Key words: , systematics, barcoding, mitochondrial DNA, short-range endemism, , Plesiolena

Introduction

The Actinopodidae Simon, 1892 is a small family of mygalomorph spiders with a Gondwanan distribution that includes three genera: Actinopus Perty, 1833 (27 species), Missulena Walckenaer, 1805 (11 species) and Plesiolena Goloboff & Platnick, 1987 (two species). Actinopus and Plesiolena are known only from South and Central America (Platnick 2012). In contrast, Missulena Walckenaer, 1805 includes 10 species from Australia and one species, M. tussulena Goloboff, 1994, from Chile. Australian species of Missulena, commonly known as Mouse Spiders, are medium-sized spiders with a steeply elevated cephalic region and a wide eye group (Fig. 1A). Some species have received public attention due to the toxicity of their venom that has been found to be biochemically similar to that of Australian Funnel-web Spiders (family Simon, 1892) (Isbister 2004; Herzig et al. 2008; Rash et al. 2000). Missulena insignis (O.P.- Cambridge, 1877), M. occatoria Walckenaer, 1805 and M. reflexa Rainbow & Pulleine, 1918 are known for conspicuous red fangs and cephalic areas in males, although many described species, e.g. M. bradleyi Rainbow, 1914, M. dipsaca Faulder, 1995, M. granulosa (O. Pickard-Cambridge, 1869), M. rutraspina Faulder, 1995 and M. torbayensis Main, 1996 are uniformly dark brown or black in colour (Main 1956, 1996). The taxonomy of Australian Missulena was first reviewed by Womersley (1943), who recognised and diagnosed six species; only four of these are known from both male and female specimens. Main (1985) summarised the current knowledge of Missulena and proposed several taxonomic changes. Shortly after, Faulder (1995) published descriptions of two widespread new species of Missulena: M. dipsaca and M. rutraspina. More recently, Main (1996) named M. torbayensis from southwestern Western Australia and noted that “it is apparent from my field observations and an abundance of specimens (mainly males) in museums that there are many undescribed species.” Recent large-scale biological surveys in the Carnarvon Basin (Main et al. 2000) and Pilbara region (Durrant et al. 2010) of Western Australia have added substantial material of Missulena, in particular males, to the collection of the Western Australian Museum (Fig. 2). Our work is based on this material and has two aims: first, we describe males of two morphologically distinct species of Missulena from the Pilbara region. Second, we conduct a

Accepted by R. Raven: 11 Feb. 2013; published: 15 Apr. 2013 521 molecular analyses using a fragment of the cytochrome oxidase c subunit I (COI) gene from a selection of Missulena specimens from Western Australia to provide an additional and independent line of evidence in the taxonomic process, and to review the suitability of this marker for single-gene barcoding of Missulena species.

Material and methods

Morphology. Specimens used for morphological examination were preserved in 75% ethanol and examined using a Leica M80 stereomicroscope. Digital images were taken using a Leica DFC295 digital camera attached to a Leica M205C stereomicroscope controlled by the Leica Application Suite Version 2.5. Distribution maps were produced using the Open Source software package Quantum GIS Version 1.7.4 Wroclaw (www.qgis.org; accessed 30 May 2012). Climate data for the Pilbara bioregion were retrieved from the Australian Government, Bureau of Meteorology (online under www.bom.gov.au/climate/averages/tables) and refer to the airport of Newman. Measurements are expressed in millimetres and were taken in dorsal view, except those of labium, sternum, coxae, trochanters and spigots, which were taken in ventral view. Total lengths were taken in dorsal view and exclude the chelicerae and spinnerets. The number of teeth on the claws is given as the formula “leg number: number of teeth of lateral claws / number of teeth of median claw”. The leg formula is given as the order of the leg lengths from longest to shortest. The leg “index” is given here as the leg length divided by carapace length and indicates the ratio of leg lengths versus carapace. Measurements of the eye region refer to the distance between the two most separate eyes in a row; those of the ocular quadrangle (OQ) to the quadrangle limited by the four lateral eyes, and those of the median ocular quadrangle (MOQ) to the distance between the four median eyes. The term “rasps” refers to the presence of short but strong conical spines at the patellae of legs I–IV. The occurrence of such spines on patella I has been suggested as a potential synapomorphy for Missulena species (Goloboff & Platnick 1987). Our species hypotheses are based only on males, as are those established by previous authors (Womersley 1943, Faulder 1995, Main 1996). All specimens are lodged at the WA Museum (WAM). Molecular methods. We sequenced 22 specimens of Missulena from Western Australia (Figs 2A, B, 3; Table 1). Reference specimens were from five additional bioregions of Western Australia (Thackway & Cresswell 1995): Avon Wheatbelt, Jarrah Forest, Gascoyne, Murchison and Ord Victoria Plain (Figs 2A, 3). Each specimen sequenced as part of this study is denoted by a superscript DNA code in the Material Examined lists, following Harvey et al. (2012). We also sequenced two putatively undescribed species of Conothele Thorell, 1878 (Ctenizidae Thorell, 1887) and retrieved an additional sequence of Euagrus chisoseus Gertsch, 1939 (Dipluridae Simon, 1889) from Genbank for use as an ultimate outgroup (Table 1). All specimens were sequenced for variation at the mitochondrial cytochrome c oxidase subunit I (COI) gene which is preferred here over other potential barcoding genes because of the general lack of introns and infrequency of nucleotide polymorphisms, high inter- and intra-specific variability, and unproblematic amplification using standard methodology (Folmer et al. 1994, Hebert et al. 2003). Genomic DNA was extracted from leg muscle tissue using the QIAGEN DNeasy Blood and Tissue Kit for animal tissues (QIAGEN Inc., Valencia, CA). Polymerase chain reaction (PCR) amplification of an 827 bp fragment of the COI gene was achieved using the primers LCO1490 (Folmer et al. 1994) and HCOoutout (Prendini et al. 2005). PCR products were sequenced for both DNA strands at the AGRF node in Perth (Western Australian Institute of Medical Research, WA, Australia). Sequences were checked by eye, edited and aligned using the program Geneious Version 5.6 (Drummond et al. 2011). Tree building. Two Bayesian phylogenetic analyses were executed using MrBayes Version 3.2.1 for Macintosh (Huelsenbeck & Ronquist 2001; Ronquist & Huelsenbeck 2003) applying 1) three partitions according to codon position, and 2) one partition across all data. MrModeltest Version 3.7 (Posada & Crandall 1998) was used to estimate the appropriate model of nucleotide substitution under an Akaike Information Criterion (AIC) framework. A general time reversible model (GTR) (Rodríguez et al. 1990) with a class of invariable sites (I) and gamma distributed rate heterogeneity (G) (Yang 1996) was invoked for the unpartitioned data and codon positions 1 and 2. A GTR model with gamma distributed rate heterogeneity (GTR+G) was suggested for the third codon position. Four Markov chain Monte Carlo (MCMC) chains were run for 40,000,000 generations to achieve convergence of the independent runs, sampling every 1,000 generations and discarding the first 25% of sampled

522 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU trees as ‘burnin’. FigTree Version 1.3.1 (Rambaut 2009) was used to visualise and edit tree files, and Tracer Version 1.5 (Rambaut & Drummond 2009) was used to check parameter values and convergence of the MCMC chains. To infer the degree of genetic similarity between the sequenced specimens, we finally calculated a distance matrix in Geneious for the sequenced Missulena specimens using the edited and aligned DNA sequence data.

Abbreviations AME = anterior median eyes, ALE = anterior lateral eyes, MOQ = median ocular quadrangle, OC = ocular quadrangle, PME = posterior median eyes, PLE = posterior lateral eyes, PMS = posterior median spinnerets, PLS = posterior lateral spinnerets, d = dorsal, pv = proventral, rv = retroventral, v =ventral.

FIGURES 1A−B. A, Missulena langlandsi sp. nov., holotype male (WAM T115948); B, floodplain of Weeli Wolli Creek in the Pilbara region of Western Australia, type locality of M. langlandsi sp. nov.

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 523 (PT). in Western Australia) in Australia) Western GenBank accession number E. chisoseus cation (all except oordinates (WGS84) Lo ographic c ographic ) Registration number Ge number ) Registration NA NA NA NA FJ607564 M T60398 WAM 118°33’00”E 31°45’00”S; Road Stirrat Muntadgin, KC708094 (PT) (PT) M T112076 WAM 119°03’52”E 23°07’09”S; Newman of WNW Wonmunna, KC708083 (PT) (PT) M T91914 WAM 119°02’50”E 22°55’54”S; Creek Wolli Weeli Range, Hamersley KC708082 (PT) (PT) M T91911 WAM 119°02’16”E 22°55’33”S; Creek Wolli Weeli Range, Hamersley KC708081 (HT) (HT) M T97017 WAM 119°16’02”E 22°58’04”S; Newman of NW 60 km Jinayri, KC708092 -group -group M T62264 WAM 120°24’36"E 22°39’16”S; Station Hill Roy E of Mt Lewin, SE of KC708087 M T95775 WAM 127°59’07”E 17°39’42”S; Mine Nickel Kimberley Malay Sally KC708096 sp. nov. sp. nov. sp. nov. ruinosa sp. nov. sp. sp. F T112337 WAM 118°59’34”E 23°09’34”S; (Pilbara) Newman of WNW Wonmunna, KC708091 sp. sp. sp. M sp. M T96437 WAM M 120°27’41”E 27°27’26”S; T91631 WAM Leinster of NNW Downs, Albion 117°11’00”E 31°08’00”S; T104384 WAM Reserve Nature Minnivale Minnivale, 116°03’33”E 32°11’42”S; Dam Wungong Reservoir, Wungong KC708085 KC708088 KC708095 Specimens sequenced for the molecular phylogenetic analyses. Sequenced type specimens are denoted as holotype (HT) or paratype holotype as are denoted specimens type Sequenced analyses. phylogenetic molecular the for sequenced Specimens sp. sp. F T112453 WAM 120°34’37”E 27°15’18”S; (Murchison) Leinster of N 50 km KC708090 sp. sp. sp. F J T95395 WAM T118178 WAM 120°15’27”E 23°22’44”S; 117°29’49"E 22°46’01”S; Newman E of 35 km minesite, Jimblebar Price Tom of east Station, Rocklea KC708074 KC708093 sp. sp. sp. sp. M M M T96335 WAM T96357 WAM 120°27’41”E 27°27’26”S; T96442 WAM 120°27’40”E 27°27’29”S; Leinster of NNW Downs, Albion 120°27’40”E 27°27’29”S; Leinster of NNW Downs, Albion Leinster of NNW Downs, Albion KC708089 KC708084 KC708086 sp. sp. sp. sp. sp. sp. F sp. F sp. F M J J T97637 WAM J T99600 WAM T102165 WAM 118°30’55”E 22°07’40”S; T113598 WAM 118°30’14”E 22°52’30”S; 120°26’48”E 23°25’44”S; T113626 WAM Station Downs Mulga Hills, Murray 119°08’21”E 23°00’11”S; T113653 WAM Downs to Juna Siding Bellbird 118°55’06”E 22°39’23”S; T113660 WAM Newman of W Creek, Davidson 118°52’50”E 22°44’33”S; Newman of NW Flank, Southern 118°52’12”E 22°38’01”S; Newman of NW 114 km Newman of NW 110 km KC708097 Newman of NW 119 km KC708076 KC708075 KC708077 KC708079 KC708078 KC708080 Missulena Missulena M. langlandsi M. langlandsi M. langlandsi Ctenizidae Conothele Conothele Pilbara faulderi M. Taxon Taxon Dipluridae chisoseus Euagrus Actinopodidae (J = juvenile Sex Missulena p Murchison Missulena Missulena Jarrah Forest Missulena Missulena Missulena Missulena Avon Wheatbelt granulosa Missulena Missulena Missulena Missulena Missulena Missulena Missulena Missulena (Kimberleys) Plain Victoria Ord TABLE 1.

524 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU Systematics

Family Actinopodidae Simon, 1892

Genus Missulena Walckenaer, 1805

Missulena Walckenaer, 1805: 8. Type species: M. occatoria Walckenaer, 1805, by monotypy. Eriodon Latreille, 1804: 134 (nomen nudum, see Pickard-Cambridge 1903). Type species: Eriodon occatorius Latreille, 1806, by monotypy (also see Judson 2012: 1275).

Missulena faulderi sp. nov. Figs 2, 4–7, 12A, B, C

Type material. AUSTRALIA: Western Australia: holotype male, Jinayri, ca. 60 km NW of Newman, 22°58’04”S, 119°16’02”E, 1 February 2009, pitfall trap, J. Gollan, N. Sullivan, M. Semeniuk & M. Beatson (WAM T97017DNA). Paratypes: 1 male, same data as holotype, except 23°02’27”S, 119°16’47”E, 2 February 2009 (WAM T96133); 1 male, as for holotype, except 23°02’30”S, 119º16’40”E, 5 February 2009 (WAM T96132). Etymology. The specific epithet is a patronym in honour of Richard Faulder for his work on Missulena (Faulder 1995).

FIGURES 2A–B. Distribution records of Missulena spp. in Western Australia: A, records of Missulena compiled from the WA Museum database (grey circles) and locality data for specimens included in molecular analyses (red circles). The biogeographic (IBRA) regions (Thackway & Cresswell 1995) are displayed in yellow. Bioregions from which specimens were sequenced are shaded darker. B, close-up of the Pilbara bioregion in Western Australia, highlighting the distribution of Missulena faulderi sp. nov. (green squares) and M. langlandsi sp. nov. (blue triangles).

Diagnosis. Males of M. faulderi sp. nov. differ from all other Missulena from Australia by the very long and slightly curved embolus of the male bulb. They differ from other Missulena, except the most similar species M. rutraspina, by the presence of a pronounced rastellum, which is developed as a distinct process with three to five large conical spines rather than a series of narrow spines on a process or low mound. Missulena faulderi sp. nov. males (carapace length < 3.5) differ from the males of M. rutraspina (carapace length > 3.8) by the smaller size, the presence of an interior row of seven to eight fused teeth in the cheliceral furrow (teeth divided in M. rutraspina), the shape of the male bulb which has no depressions, and a medially curved embolus (bulb with spiral depression and embolus almost straight in M. rutraspina, see Faulder 1995). Missulena faulderi sp. nov. males do not have any red markings on the carapace and chelicerae (as do males of M. occatoria and M. langlandsi sp. nov.), do not

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 525 have rasps on patellae I, II (similar to M. dipsaca, M. rutraspina & M. tussulina) and IV, and have lateral ridges on the chelicerae. Females of M. faulderi sp. nov. are unknown.

FIGURE 3. Bayesian majority-rule consensus tree showing results from a partitioned phylogenetic analysis of the COI mtDNA dataset (25 taxa, 826 bp). Species described in this study are colour-coded in orange. The three major uncovered lineages are shaded light blue, green and red. Posterior probabilities are calculated in %. Bioregions (see Fig. 2A): AW – Avon Wheatbelt, GAS – Gascoyne, OVP – Ord Victoria Plain, JAR – Jarrah Forest, MUR – Murchison.

Description. Adult male, based on holotype WAM T97017. Medium-sized (total length 7.5). Colour: carapace (Figs 4A–C) dark reddish-brown but distally lighter, margins dark brown; eye region (Fig. 5D) dark brown, anterior median eyes on black tubercle; chelicerae (Figs 4B, 5B, E) proximally dark brown but distally lighter, fangs (Figs 5A, C) dark brown; abdomen (Figs 4A, D) dark grey with yellow longitudinal streaks and two proximomedial pale patches of two to three individual spots each, mottled with some more spots posteriorly which form oblique lines; sternum (Fig. 6A) yellowish-brown, margins contoured greyish-brown, sigilla orange-brown; labium (Figs 6B, D) and maxillae dark reddish-brown; legs (Figs 4A, E, F) yellowish grey- brown, tarsi and metatarsi ventrally yellow; spinnerets (Fig. 6C) pale yellow, spigots white. Carapace: 3.5 long, 3.9 wide; caput and eye region (Fig. 4C) elevated in lateral view, strongly arched and differentiated from lower carapace; fovea (Fig. 4G) very deep, strongly procurved (Fig. 4B); lower carapace rugose with bands of fine, random fissures centered around fovea (Fig. 4B).

526 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU FIGURES 4A−G. Missulena faulderi sp. nov., holotype male (WAM T97017): A, habitus, dorsal view; B, carapace, dorsal view; C, same, lateral view; D, abdomen, dorsal view; E, patella III, dorsal view; F, patella II, dorsal view; G, fovea, dorsal view.

Eyes: OQ 3.4 times wider than long, occupies 0.76 of cephalic width; MOQ 1.42 long; width of anterior eye group 2.3, width of posterior group 1.8, OQ length 0.6; AME on tubercle, 0.6 wide, 0.4 long; AME inter-distance 0.1; AME to ALE 0.35; PLE to ALE 0.4; PME to ALE 0.25; PLE to PME 0.35; eye region (Fig. 5D) with reduced setation but some setae anterior to AME. Chelicerae: 2.1 long, 1.3 wide; distally broad, diagonal, slightly conical; edges smoothly rounded; with very strong transverse ridges distally over entire length (Figs 5B, E), without setae in area of transverse ridges but with 10 setae along inner margin of chelicera; rastellum (Figs 5A, C) strongly developed, very pronounced, consisting of sclerotised process with four strong conical spines and 20−22 disordered setae (Figs 5A, E), 10 long setae extend forward from anterior margin of each chelicera and cover base of fang, setae largest on latero-ventral side; inner margin of cheliceral furrow with three rows of teeth (Figs 5F, G); prolateral (inner) row with 13 teeth, distal teeth

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 527 entirely fused and forming blade, all teeth fused at base; intermediate row with four proximal, spaced teeth; retrolateral (outer) row with two proximal, spaced teeth; without basomedial teeth.

FIGURES 5A−G. Missulena faulderi sp. nov., holotype male (WAM T97017): A, tip of chelicerae, white rectangle indicates rastellum; B, chelicerae and eye group, frontodorsal view; C, rastellum, ventral view; D, eye group, dorsal view; E chelicerae, frontal view; F, detail of right cheliceral groove, inner row of teeth; G, same, depicting inner, medial and outer row of teeth.

Maxillae: 1.6 long; 1.1 wide, almost square (Fig. 6E), 50 pointed cuspules along entire anterior margin, distally pointed and extended onto prominent heel. Labium: 0.7 long, 0.65 wide; conical, 18 pointed cuspules anteriorly (Fig. 6D); labiosternal suture developed as shallow groove; pair of sigilla near labiosternal suture (Fig. 6A), developed as irregular, poorly-defined patches. Sternum: 2.2 long, 1.8 wide; oval, rebordered (Fig. 6A); with prominent setae, arranged irregular but denser lateral to labium; three pairs of sigilla, anterior pair smallest but well defined, posterior pair roughly oval but not well defined, all sigilla slightly depressed.

528 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU FIGURES 6A−E. Missulena faulderi sp. nov., holotype male (WAM T97017): A, sternum, ventral view; B, labium and chelicerae, ventral view; C, spinnerets, ventral view (paratype WAM T91911); D, labium, ventral view; E, right maxilla, ventral view.

Abdomen: 3.7 long, 3.0 wide; roughly oval, without bumps or processes (Fig. 4D); four spinnerets (Fig. 6C), PLS 0.9 long, 0.4 wide; PMS 0.42 long, 0.2 wide. Pedipalp: length of trochanter 0.7, femur 2.3, patella 1.2, tibia 1.8, tarsus 0.5; entire palp aspinose, tarsus terminally blunt (Figs 7B, C); bulb pyriform and more stout than globular (Figs 7A, B, F, H), two strongly sclerotised sections connected by median haematodocha (Fig. 12C); bulb strongly twisted proventrally (Figs 7A, G); embolus very long, tapering and slightly twisted medially (Figs 12A, B, C); embolus tip simple, triangular, without processes (Figs 7D, 12A, B, C). Legs: with less than 150 brown setae over all segments, ventral setae of tibiae and metatarsi generally much longer and thicker than dorsal setae and bent towards exterior, dorsal and lateral setae of tibiae and metatarsi shorter than diameter of respective segment, ventral setae as long or longer than respective leg segment; preening comb absent (Figs 4A, E, F); metatarsi I and II ascopulate, tarsi I ascopulate but with 80 fine ventral setae distally, apical sections with tiny ventral scopula; tarsi II slightly scopulate ventrally across entire length but fine setae of scopula not very dense; tarsi and metatarsi III and IV scopulate ventrally across entire length. Leg spination: pedipalp aspinose; leg I: tibia rv0, v12, pv0, d0; metatarsus rv3, v17, pv0, d0; tarsus rv3, v4, pv2, d0; leg II: tibia rv0, v12, pv0, d0; metatarsus rv4, v8, pv0, d0; tarsus rv4, v4, pv3, d0; leg III: tibia rv0, v16, pv8, d0; metasarsus rv12, v0, pv7, d11; tarsus rv3, v3 (apical), pv3, d3 (apical); leg IV: tibia rv 13, v0, pv0, d0; metatarus rv10, v0, pv2, d0; tarsus rv12, v7 (apical), pv5, d2 (apical); patellae I, II without rasps and spines, patellae III with 31 rasps in eight oblique rows dorsally, median rows shorter than lateral rows and with less spines (Fig. 4E); patella four with three to four thick, short spines (Fig. 4F) retrolaterally.

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 529 FIGURES 7A−H. Missulena faulderi sp. nov., holotype male (WAM T97017): A, bulb and tarsus, dorsal view; B, left pedipalp, retroventral view; C, left pedipalp, proventral view; D, tip of embolus, dorsal view; E, bulb and embolus, dorsolateral view; F, same, dorsal view; G, tibia and bulb, retroventral view; H, same, proventral view.

Leg measurements: leg formula IV>I>II>III. Leg I: coxa 1.4, trochanter 0.8, femur 2.8, patella 1.4, tibia 2.0, metatarsus 1.8, tarsus 1.1, total = 11.3; tibia width 0.7, leg I index = 3.2. Leg II: coxa 1.3, trochanter 0.9, femur 2.6, patella 1.4, tibia 2.1, metatarsus 1.6, tarsus 1.0, total =10.9; tibia width 0.8, leg II index = 3.1. Leg III: coxa 1.4, trochanter 0.7, femur 2.3, patella 1.3, tibia 1.4, metatarsus 2.1, tarsus 1.3, total = 10.5; tibia width 0.7, leg III index = 3.0. Leg IV: coxa 1.4, trochanter 0.8, femur 3.4, patella 1.3, tibia 2.2, metatarsus 2.3, tarsus 1.3, total = 12.7; tibia width 0.7, leg IV index = 3.6. Trichobothria: arranged in discontinuous rows; tibiae I–IV with two rows of 3−4 in dorsolateral position, first row prodorsal and second row retrodorsal, metatarsi with 4 mediodorsal, tarsi I+II with 3, III+IV with 4 mediodorsal.

530 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU Tarsal claws: leg I: 5−6/ 2; leg II: 3−4/ 1; leg III: 1−2/ 1; leg IV: 1−2/ 0−1, teeth on claws III−IV very small and broad; claws slightly shorter than spines of tarsi. Variation in paratypes (N=2): total length 6.3–7.8; carapace 3.4–3.7 long, 3.9–4.0 wide; number of labial cuspules 9–13, maxillary cuspules 28–38; labial groove may be inconspicuous; rastellum with 3–5 thick, conical spines. Distribution. Currently known only from the Jinayri exploration lease (BHP Billiton Iron Ore) in the Pilbara region northeast of Newman (Fig. 2B). Phenology and habitat preferences. All males were collected in pitfall traps in February, the month of highest annual rainfall in the Pilbara region. The holotype was collected adjacent to a then dry creek bed near Weeli Wolli Creek.

FIGURES 8A−H. Missulena langlandsi sp. nov., holotype male (WAM T115948): A, habitus, dorsal view; B, carapace, dorsal view; C, carapace, lateral view; D, fovea, dorsal view; E−F, patella I, dorsal view; G, patella III, dorsal view; H, patella IV, dorsal view.

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 531 Missulena langlandsi sp. nov. Figs 1, 2, 8−11, 12D, E, F

Type material. AUSTRALIA: Western Australia: holotype male, Wonmunna mining lease, ca. 73 km heading 291° from Newman, 23°07’16”S, 119°03’48”E, 20 May 2011, hand-collecting, P. R. Langlands (WAM T115948). Paratypes: 1 male, same data as holotype except 23°07’09”S, 119°03’52”E, 20 May–22 June 2011, wet pitfall trap (WAM T112076 DNA) Other material examined. AUSTRALIA: Western Australia: 1 male, Hamersley Range, Weeli Wolli Creek region, 22°55’54”S, 119°02’50”E, 17 June 2008, P. Bolton & J. Puglisi (WAM T91914 DNA); 1 male, same data except 22°55’53”S, 119°02’49”E (WAM T91912); 1 male, same data except 22°55’53”S, 119°02’50”E (WAM T91913); 1 male, same data except 22°55’33”S, 119°02’16”E, 15 June 2008 (WAM T91911 DNA). Etymology. The specific epithet is a patronym in honour of Peter Langlands, the collector of the holotype, for his keen interest in natural history and nature conservation. Diagnosis. Males of M. langlandsi sp. nov. differ from other red-headed Missulena (M. occatoria, M. insignis and M. reflexa) in their small body size (about < 5.0 in M. langlandsi, > 8.0 in other species), small number of cheliceral teeth (retrolateral row with less than 7 teeth), rastellum with less than 8 spines, patellae III with less than 30 rasps, patellae IV with less than 15 rasps, and the undivided second pair of sigilla (see Womersley 1943: 262). Females of M. langlandsi sp. nov. are unknown. Description. Adult male, based on holotype WAM T115948. Small spider (total length 4.9). Colour: carapace base colour black, caput and chelicerae uniformly orange-red (Figs 8A–D); eye region orange-red but ALE, AME and PLE on black tubercles (Fig. 8B); chelicerae with black setae, fangs orange-brown; abdomen greyish-brown with black setae, distal portion with light brown transverse lines which are diagonally oblique (Figs 8A, 10G); sternum distally olive-green but proximally orange, margins contoured greyish-orange (Fig. 9A), sigilla orange-brown, many grey setae; labium uniformly orange, maxillae olive-green but heel orange (Fig. 9B); legs black or dark grey, articles greyish-white (Fig. 8A); spinnerets greyish-brown, spigots brown (Fig. 9C). Carapace: 2.4 long, 2.9 wide; caput and eye region elevated in lateral view, strongly arched and differentiated from lower carapace (Fig. 8C); fovea deep, strongly procurved (Fig. 8D); lower carapace rugose with bands of fine, random fissures centered around fovea (Fig. 8B). Eyes: OQ 4.3 times wider than long, occupies 0.72 of cephalic width; MOQ 1.15 long; width of anterior eye group 1.9, width of posterior group 1.7, OQ length 0.43; AME on tubercle, 0.4 wide, 0.25 long; AME inter- distance 0.08; AME to ALE 0.35; PLE to ALE 0.25; PME to ALE 0.25; PLE to PME 0.2; eye region with black setae, one straight line of ca. 10 setae between AME and fovea; two other oblique lines of 10 setae between fovea and lateral eyes (Fig. 10D). Chelicerae: 1.7 long, 1.0 wide; distally broad and diagonal, slightly conical; edges smoothly rounded; outer surface smooth and without ridges (Fig. 10E); proximal parts without setae, distal and interior parts with ca. 70 setae, these setae distally increase in size (Figs 10A–C); rastellum not very pronounced, a shallow process with 7−10 basally thickened black setae interspersed with 9−12 longer and thinner setae, thick setae ca. 1/3 shorter and thicker than others; 10−12 long setae extend forward from anterior margin of each chelicera and cover base of fang, setae largest on latero-ventral side; inner margin of cheliceral furrow with two rows of teeth, prolateral (inner) row consists of 7 spaced, conical teeth, retrolateral (outer) row with two proximal, spaced teeth; basomedial tooth present (Figs 10E, F). Maxillae: 1.3 long; 0.8 wide; almost square, 22–25 small cuspules along entire anterior margin but distally sparser (Fig. 9E). Labium: 0.7 long, 0.6 wide; conical, ca. 8–10 small cuspules distally (Fig. 9D); labiosternal suture as shallow groove (Fig. 9B); pair of sigilla near labiosternal suture, kidney-shaped and well-defined. Sternum: 1.7 long; 1.6 wide; oval, rebordered; many long setae present, three pairs of sigilla, anterior two pairs smaller than most posterior pair, all well defined, depressed and roughly oval (Fig. 9A). Abdomen: 2.4 long, 2.8 wide; roughly oval without bumps or processes (Fig. 10G); four spinnerets, PLS 0.9 long, 0.4 wide, apical segment domed (Fig. 9C); PMS 0.40 long, 0.2 wide.

532 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU FIGURES 9A−E. Missulena langlandsi sp. nov., holotype male (WAM T115948): A, sternum, ventral view; B, labium and chelicerae, ventral view; C, spinnerets, ventral view; D, labium, ventral view; E, right maxilla, ventral view.

Pedipalp: length of trochanter 0.4, femur 2.2, patella 1.1, tibia 2.2, tarsus 0.9; entire palp aspinose, tarsus terminally blunt (Figs 11D, E); bulb pyriform (Figs 11A, B, D, E), two strongly sclerotised sections connected by median haematodocha (Fig. 12F), bulb slightly twisted proventrally; embolus medium-sized, tapering and slightly bent medially, directed upwards (Figs 12D, E, F); embolus tip simple, without processes (Fig. 11C). Legs: covered uniformly with more than 250 long and thin black setae; setae of tibiae, metatarsi and tarsi as long or longer than diameter of respective segment; preening comb absent; legs I–II ascopulate, metatarsi III scopulate ventrally across distal half, metatarsi IV and tarsi III–IV ventrally scopulate across entire length. Leg spination: pedipalp aspinose; leg I: tibia rv0, v13, pv0, d0; pv0, d0; tarsus rv0, v4, pv0, d0; leg II: tibia rv0, v13, pv0, d0; metatarsus rv0, v6, pv0, d0; tarsus rv0, v5, pv2, d0; leg III: tibia rv5, v13, pv3 (apical), d0; metatarsus rv5, v6, pv3, d6; tarsus rv1, v9, pv4, d2 (apical); leg IV: tibia rv2 (apical), v6, pv0, d0; metatarsus rv8, v11, pv0, d1 (apical); tarsus rv9, v19, pv0, d2 (apical); patellae I with two apical spines (Figs 8E, F); patellae II with three apical spines dorsally; patellae III with 22 rasps in five discontinuous rows dorsally (Fig. 8G); patellae IV with five irregular spines dorsally (Fig. 8H). Leg measurements: leg formula IV>I>II>III. Leg I: coxa 1.2, trochanter 0.7, femur 2.5, patella 1.25, tibia 1.6, metatarsus 2.0, tarsus 1.1, total = 10.35; tibia width 0.5, leg I index = 4.3. Leg II: coxa 1.2, trochanter 0.7, femur 2.3, patella 1.25, tibia 1.85, metatarsus 1.8, tarsus 1.2, total = 10.3, tibia width 0.6, leg II index = 4.3; Leg III: coxa 1.2, trochanter 0.7, femur 2.3, patella 1.1, tibia 1.4, metatarsus 1.7, tarsus 1.2, total = 9.6; tibia width 0.7, leg III index = 4.0. Leg IV: coxa 1.2, trochanter 0.6, femur 3.4, patella 1.3, tibia 1.8, metatarsus 1.9, tarsus 1.2, total = 11.4; tibia width 0.65, leg IV index = 4.75.

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 533 FIGURES 10A−G. Missulena langlandsi sp. nov., holotype male (WAM T115948): A, fangs, frontoventral view; B, chelicerae, frontal view; C, tip of chelicerae, white rectangle indicates that rastellum is poorly developed; D, eye group, dorsal view; E−F, right cheliceral groove, depicting inner and outer row of teeth; G, abdomen, dorsal view.

Trichobothria: arranged in discontinuous rows; tibiae I–IV with two rows of 3 in dorsolateral position, first row prodorsal and second row retrodorsal; metatarsi with 4 mediodorsal, tarsi I–II with 3 and tarsi III–IV with 4 mediodorsal. Tarsal claws: leg I: 5−8/ 3; leg II: 4−5/ 2; leg III: 4−5/ 1; leg IV: 0−1/ 1, all teeth prominent and clearly divided; claws about as long as spines of tarsi. Variation in paratypes (N=5): total length 4.6–5.2, carapace 2.4–2.8 long, 2.9–3.1 wide; cephalic region may be slightly granulate; prolateral row of cheliceral teeth consists of 4–7 teeth; one or two more teeth may be present basally in fang groove between the two rows of teeth; labial cuspules may be absent or reduced to a few tiny grey spots, maxillary cuspules may be small and present as 15−30 tiny grey spots.

534 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU The spination of the rastellum varies considerably. One paratype (WAM T112076) has three thick spines in addition to the basally thickened setae on the right, but not the left chelicera. A third specimen (WAM T91911) has three to four thick spines on a prominent elevation but no basally thickened setae. The character is variable between and even within specimens and care must be taken in using it for the identification of this species. The rastellum is slightly more elevated and the number of basally thickened setae is lower in specimens with prominent spines whereas specimens without spines have generally more thickened setae on a very shallow elevation.

FIGURES 11A−E. Missulena langlandsi sp. nov., holotype male (WAM T115948): A, left pedipalp, retroventral view; B, same, proventral view; C, tip of embolus, dorsal view; D, bulb and embolus, proventral view; E, same, retroventral view.

Distribution. Missulena langlandsi sp. nov. is currently only known from the floodplain of Weeli Wolli Creek north-west of Newman (Fig. 2B). Phenology and habitat preferences. Males of M. langlandsi sp. nov. were collected from May to June when annual average temperatures in the Pilbara are at their lowest and rainfall decreases. The holotype of M. langlandsi sp. nov. was collected adjacent to the then dry creek bed of Weeli Wolli Creek in open habitat dominated by spinifex (Triodia spp.) and introduced Buffel grass (Cenchrus ciliaris) with scattered gum trees (Eucalyptus spp./ Carymbia spp.) (Fig. 1B). It was collected during the day, wandering around in bright sunshine.

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 535 FIGURES 12A−F. Male bulb and embolus of Missulena faulderi sp. nov., holotype male (WAM T97017) and Missulena langlandsi sp. nov., holotype male (WAM T115948). Missulena faulderi sp. nov.: A, prolateral view; B, same, ventral view (slightly twisted retrolaterally) ; C, same, dorsal view. Missulena langlandsi sp. nov.: D, ventral view; E, same, dorsal view (slightly twisted retrolaterally); F, same, prolateral view.

Results and Discussion

Implications of molecular analyses. Partitioned and unpartitioned Bayesian analyses resulted in identical tree topologies and suggested three major lineages of Missulena in Western Australia within our dataset (Fig. 3): lineage I consisted of three single-specimen branches and included the holotype of M. faulderi sp. nov., lineage II included three putative species and lineage III included five putative species, all represented by red-headed forms of Missulena, including the new species M. langlandsi sp. nov. All lineages and clades received strong statistical support (posterior probability values >90%) in the analyses, except clade IIIc, which received low support values (69%) in the partitioned analysis.

536 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU The cladograms from the morphological and molecular data conflict; this is perhaps highlighted best by the exploration of sequence divergences between the specimens we have attributed to the clade M. langlandsi sp. nov. and those forming clade IIIb (Fig. 3). The specimens belonging to clade IIIb differ from M. langlandsi sp. nov. in overall body size (total length ca. 9–10 versus 5.0 in P. langlandsi sp. nov.) but also in a suite of key- taxonomic characters which, when treated in their entirety, do not allow us to refer to both clades as the same species. Yet, these clades are strongly affiliated in the phylogenetic analyses and form a monophyletic group with maximum statistical support. Genetic divergences between both clades are less than 9% (Table 2), and based solely on the COI dataset, we could have hypothesised that both clades belong to a common species, which is in sharp contrast to the morphological implications. There are further inconsistencies between morphology and the COI sequences: two specimens identified as the same morphospecies prior to the molecular analysis and monophyletic in our topology, clade IIIc (Fig. 3), showed a sequence divergence of 12.1% (Table 2). Based on previously suggested species divergence limits for , e.g. > 5% divergence in Parmakelis et al. (2006); > 8% in Hebert et al. (2003), these should be considered separate species. In a subsequent re- examination of the respective specimens we were, however, unable to find morphological characters to justify such an approach, indicating either homoplasy in the morphological characters or lack of resolution in the single molecular character used – the COI sequence data. The second hypothesis is clearly more parsimonious because it does not require homoplasy and/or multiple transformations in a whole range of putatively unlinked morphological characters whereas COI can be treated as a single functional unit in which the characters (i.e. the individual base pairs) cannot evolve freely due to functional constrains. Thus, COI alone fails to clearly resolve Missulena species and is most likely of limited value in barcoding these spiders. We recommend that additional independent genetic markers be used to delineate species of Missulena, with nuclear protein-coding genes being potential candidates in a preferably multi-gene approach towards species delineation (Bailey et al. 2010, Blackledge et al. 2009, Bond & Stockman 2008, Satler et al. 2011). Morphological characters. Some morphological characters in M. faulderi sp. nov. highlight the importance of this species for future phylogenetic studies on Missulena. The first is the presence of a strongly- sclerotised rastellum on a distinct process in M. faulderi sp. nov. and M. rutraspina. Such a process is absent in other known species of Missulena but also in Plesiolena, both of which are characterised by having a rastellum on a low mound (Raven 1985, Goloboff & Platnick 1987). The presence of a distinct process in M. faulderi sp. nov. and M. rutraspina is, however, shared with the South-American genus Actinopus that is characterised by having a distinct rastellum on a long projection (Raven 1985). Additional characters distinguish M. faulderi sp. nov. from other described species. Although it has toothed claws like other species from Australia (Womersley 1943), these are not as long and pronounced as in the red-headed species, e.g. M. langlandsi sp. nov., which is important because this character appears to be of phylogenetic significance (Raven 1985, Goloboff & Platnick 1987). The anterior sigilla are not small and oval as in other Missulena species (Raven 1985) but rather indistinct and consist of several small patches. The leg spines are also significantly shorter and thicker than in red-headed species from Australia. These characters are clearly important but need to be evaluated as part of a comprehensive phylogenetic analysis. Distribution and implications for assessments. The two species described here show very narrow distributions. Despite intense survey work in this resource-rich area of the Pilbara, M. faulderi sp. nov. and M. langlandsi sp. nov. are currently known only from a very confined range north-west of Newman (Fig. 2B). Both species currently classify as short- or narrow-range endemic taxa according to Harvey (2002) and Ponder and Colgan (2002), contradicting previous suggestions that many species belonging to Missulena are widespread (Faulder 1995, Main 1976). Species with wide ranges, e.g. M. occatoria from southern Australia, may in fact represent a mosaic of cryptic species with potentially short ranges and we agree with Main (1996) that “Even the type species, M. occatoria Walckenaer as it is currently defined, appears to have several other species confused with it”. It will be impossible to diagnose species of Missulena conclusively unless a detailed taxonomic study, including a suite of morphological and molecular methods, is undertaken and a comprehensive taxonomic framework has been established.

NEW SPECIES OF MOUSE SPIDERS FROM THE PILBARA REGION Zootaxa 3637 (5) © 2013 Magnolia Press · 537 M. 11.9 occatoria T62264 18.0 17.2 16.3 15.4 15.4 15.2 15.4 15.5 28.7 14.3 14.9 16.8 16.4 18.5 14.1 13.5 13.3 12.2 12.7 group

T96437 8.9 8.2 8.0 3.1 1.1 17.4 16.8 15.2 14.4 14.3 13.9 13.7 13.9 28.3 15.0 14.1 15.2 16.9 18.4 87.3

lower half the percentage of the half lower Missulena T96357 8.7 8.0 7.7 2.3 sp. 17.5 16.9 15.4 14.9 14.6 14.3 14.3 14.5 28.4 15.6 14.3 15.5 17.2 19.0 98.9 87.8

T96442 8.5 7.7 7.5 17.3 16.9 15.4 15.615.4 15.514.6 14.6 16.4 14.0 14.0 14.3 28.2 15.8 15.1 15.6 16.1 19.0 97.7 96.9 88.1

T91914 4.5 0.2 18.0 18.1 16.0 15.8 15.5 15.4 15.0 15.1 15.4 28.9 15.5 14.9 16.6 16.6 18.9 92.5 92.3 92.0 86.7 M. langlandsi T91911 4.7 18.3 18.1 16.2 16.1 15.7 15.6 15.2 15.4 15.6 29.2 15.7 15.1 16.8 16.8 19.1 99.8 92.3 92.0 91.8 86.5 sp. nov. T112076 19.0 18.4 17.0 16.3 16.0 15.7 15.5 15.5 15.6 29.4 16.0 15.6 16.4 16.8 18.5 95.3 95.5 91.5 91.3 91.1 85.9

M. T95775 pruinosa 18.1 81.5 80.9 81.1 M. T60398 granulosa 18.0 83.2 83.2 83.4 Missulena T104384 sp. 16.8 83.6 83.2 83.4

T96335 Missulena 15.7 84.4 84.9 85.1 sp. specimens are shaded grey. grey. are shaded specimens T91631 16.8 84.0 84.3 84.5

sp. nov. sp. nov. T97637* 31.7 70.6 70.8 71.1

T113660 16.7 84.4 84.4 84.6 M. langlandsi M. T113626 16.6 84.5 84.6 84.9 and and 160 bp in the alignment files are missing due to poor sequencing results. results. to poor sequencing due missing are files alignment 160 bp in the

T99600 ∼ Missulena 16.3 85.1 84.8 85.0 sp. nov. sp. T113598 17.0 84.3 84.4 84.6

T113653 16.6 84.0 84.3 84.5 specimens sequenced as part of this study. Values in upper half indicate the percentage of genetic difference and those in the in those and difference genetic of the percentage indicate half in upper Values study. this part of as sequenced specimens Missulena faulderi faulderi Missulena T95395 2.4 5.8 6.8 6.4 7.5 7.7 22.97.5 7.7 2.4 5.8 6.8 6.4 14.8 15.5 15.4 15.5 17.6 16.9 83.7 83.9 84.2

Missulena T102165 17.9 17.916.6 17.5 16.897.6 17.394.2 22.9 17.77.6 6.0 6.8 6.7 94.093.2 15.4 17.9 93.293.6 15.423.0 32.06.7 7.9 3.5 96.5 93.392.523.7 16.0 13.57.1 18.1 96.5 92.492.36.9 13.4 2.4 15.2 14.9 17.3 97.6 93.3 92.477.1 14.622.8 17.1 14.3 17.0 93.1 93.1 77.185.2 15.06.4 6.2 13.3 15.2 16.8 93.8 92.9 77.0 84.684.5 15.7 14.6 17.6 18.5 21.6 93.6 76.3 86.5 84.6 17.3 84.6 0.2 14.1 13.5 99.8 77.2 86.6 85.1 84.084.5 15.1 14.6 78.4 86.7 85.4 85.7 84.882.4 21.7 17.1 13.4 78.3 86.5 85.4 85.0 84.8 82.983.0 13.5 15.2 86.5 85.4 85.9 84.3 82.483.8 14.8 28.6 16.7 71.4 85.2 86.6 84.9 82.784.0 13.4 28.6 71.4 86.6 84.8 82.984.6 15.4 28.3 12.1 87.9 71.7 84.6 83.3 84.684.4 16.6 30.0 15.0 85.0 70.0 83.4 85.4 84.684.5 28.9 16.6 15.4 84.6 83.4 71.1 85.4 85.1 84.883.6 16.4 16.3 83.7 83.6 86.0 85.4 85.6 83.7 16.0 12.8 87.2 84.0 86.0 85.7 85.7 84.6 19.8 80.2 85.7 85.7 86.1 84.6 18.3 81.7 71.8 85.5 86.3 84.8 84.2 71.6 86.1 84.6 84.9 84.4 71.8 84.5 84.4 85.7 85.0 71.3 83.9 84.5 85.9 85.7 81.0 82.8 84.8 85.1 81.0 83.1 83.2 81.6 83.4 81.5

M. faulderi T97017

sp. nov. 16.6 83.4 83.1 83.4 83.0 83.7 83.4 83.3 69.3 83.2 84.3 83.2 82.0 81.9 81.6 81.9 81.9 83.1 83.1 83.2 82.8 Missulena T118178 sp. 83.4 81.0 81.7 82.0 distance matrix for for matrix distance COI species WA M registration T118178 T97017 82.1 T102165 82.1 T95395 82.5 T113653 83.2 T113598 82.7 T99600 82.3 T113626 82.1 T113660 68.0 T97637* 81.9 T91631 82.7 T96335 83.0 T104384 83.2 T60398 81.5 T95775 T112076 T91911 T91914 82.7 T96442 82.5 T96357 82.6 T96437 82.0 T62264 TABLE 2. TABLE first the as divergences sequence higher artificially shows T97637 WAM * identical (shared) bases between specimens. specimens. between bases (shared) identical

538 · Zootaxa 3637 (5) © 2013 Magnolia Press HARMS & FRAMENAU Acknowledgements

This research was made possible through an Endeavour International Postgraduate Research Scholarships (EIPRS) and a UPAIS top-up scholarship of the University of Western Australia to the senior author. We are thankful to the School of Animal Biology at the University of Western Australia for the use of molecular laboratory facilities. Dragon Energy Ltd, West Perth, provided funding for DNA sequencing. We thank Peter Langlands (Phoenix Environmental Sciences) and Conor O’Neill (Calibre Global) for cheerfully introducing the senior author to mygalomorph spider diversity in the field. Initial specimens of both new species were collected during invertebrate surveys on behalf of BHP Billiton Iron Ore and their support of scientific research through funding of taxonomic research positions (including for the junior author from 2009 to 2011) are greatly appreciated. We are grateful to Mark Harvey (WAM) for granting access to the collection, and Mieke Burger, Mark Castalanelli and Julianne Waldock (all WAM) for logistics support. We thank our partners, Stephanie Harms and Melissa Thomas, for their patience and support, without which this study would not have been possible. Finally, we wish to thank Pablo Goloboff and Robert Raven for providing invaluable feedback on earlier versions of this manuscript.

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