THE PHOLCIDS OF (ARANEAE; ): , BIOGEOGRAPHY, AND RELATIONSHIPS

BERNHARD A. HUBER Division of Invertebrate Zoology American Museum of Natural History present address: Institute of Zoology University of Vienna Althanstr. 14, A-1090 Wien, Austria

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 260, 144 pp., 437 ®gures, 19 maps Issued April 16, 2001 Price: $15.40 a copy

Copyright ᭧ American Museum of Natural History 2001 ISSN 0003-0090 CONTENTS Abstract ...... 3 Introduction ...... 3 Materials and Methods ...... 4 Relationships ...... 4 Biogeography ...... 7 Key to the Pholcids of Australia ...... 9 Taxonomy ...... 11 Wugigarra, new ...... 11 Trichocyclus Simon ...... 56 Micromerys Bradley ...... 95 Pholcus Walckenaer ...... 108 Panjange Deeleman-Reinhold and Deeleman ...... 118 Spermophora Hentz and Belisana Thorell ...... 124 Introduced Species ...... 132 Acknowledgments ...... 138 References ...... 138 Index of Generic and Speci®c Names ...... 140 Appendix 1. Matrix for Phylogenetic Analysis ...... 142 Appendix 2. NONA Cladogram ...... 143 Appendix 3. Pee-Wee Cladogram ...... 144

2 2001 HUBER: PHOLCIDS OF AUSTRALIA 3

ABSTRACT The pholcid of Australia are revised. Only seven autochthonous genera are recog- nized: (1) Wugigarra, new genus, includes the Australian species previously assigned to Psil- ochorus Simon. This genus is largely restricted to eastern Australia. With 22 described species (20 of them new) and about 40 undescribed species in Australian collections, it is probably the most diverse genus on the continent. (2) Trichocyclus Simon is the dominant or only pholcid genus in most areas of Western and South Australia and the Northern Territory; 23 species are described, 20 of them new. (3) Micromerys Bradley is restricted to the tropical and subtropical areas of Queensland and Northern Territory; 7 species are described, 5 of them new. (4) Pholcus Walckenaer has within Australia the same distribution as does Micromerys; 4 autochthonous species are described, all new. Pholcus litoralis Koch is newly synonymized with P. phalangioides (Fuesslin). (5) Panjange Deeleman-Reinhold and Deeleman is repre- sented by a single, previously described species in northern Queensland. (6) Spermophora is restricted to northeastern Queensland; 2 species are described, both new. (7) Belisana Thorell with a single new species in the tropical north of Queensland and Northern Territory. Nine pholcid species are introduced, some of them occurring throughout the continent. They are included in a key. A numerical cladistic analysis is performed using a matrix of 71 taxa (10 of them Australian) and 65 characters. This analysis suggests that the two highly diverse genera (Wugigarra, Trichocyclus) are most closely related to New World, African, and Middle Eastern genera. All other genera are included in the Pholcus group sensu Huber. It is argued that these are probably new tropical elements, having entered Australia from the north probably not earlier than the Pleistocene, while Wugigarra and Trichocyclus are relicts of Gondwanaland, with their pres- ence in Australia dating back to the Mesozoic.

INTRODUCTION study of these collections has never been at- tempted. Any such earlier attempt would When this project was started, it had a have been confronted with signi®cantly few- much wider scope, covering at least Austra- er specimens, because most material in the lian and southeast Asian pholcids. Of these Australian collections is quite new; that is, two areas, Australia seemed incomparably almost 70% of the specimens studied were easier to do, with just 14 species recorded, collected after 1980. ®ve of which were synanthropic cosmopoli- Thus, the history of our knowledge of the tan or cosmotropical species. However, when family in Australia is quickly outlined. The the four major loans of specimens arrived ®rst contributors were Koch (1867, 1872), from Australia, totaling about 1800 speci- Bradley (1877), and Simon (1908). Koch de- mens in 1100 vials, it soon became clear that I could either just touch the surface of the scribed one valid species (Pholcus sphaero- Australian fauna in order to keep up the orig- ides, now Wugigarra s.) and created a syn- inal scope, or rather do Australia more thor- onym of (P. litoralis); oughly and drop southeast Asia. The ques- Bradley described Micromerys gracilis, cor- tion was actually decided by the unfortunate rectly creating a new genus; and for Simon, fact that the ®rst major southeast Asian col- a couple of females of Trichocyclus nigro- lection arrived too late to be substantially in- punctatus were enough to recognize that they cluded. belonged to a new genus. These pioneers Most Australian pholcids previously de- were followed by a dormant phase of about scribed were collected by European expedi- 80 years, in which only a few records for tions or private European collectors. Time introduced species like Pholcus phalangioi- constraints and other factors probably ex- des were added (e.g., Rainbow, 1911). Fi- plain why only a few specimens of pholcids nally, a series of papers by Deeleman-Rein- were thus collected. Australian collections hold (1986b, 1993, 1995) increased the num- have been included only recently (Deeleman- ber of autochthonous species to nine, adding Reinhold, 1986b, 1993), but a more thorough Panjange as a fourth genus. 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

MATERIALS AND METHODS 67. Micromerys gracilis Bradley: Northern This work is based almost entirely on the Territory, Radon Ck. (QMB) material of four Australian institutions 68. Micromerys daviesae Deeleman-Reinhold: (AMS, QMB, SAM, WAM below). Only a Queensland, Rundle Range (QMB) few specimens (e.g., types) were studied 69. Panjange mirabilis Deeleman-Reinhold: from the other institutions listed. Further Queensland, Iron Range (QMB) Australian collections (e.g., Melbourne, Can- 70. Spermophora yao, n. sp.: Queensland, berra, Darwin) may contain important ma- Iron Range (QMB) terial but could not be incorporated because 71. Belisana australis, n. sp.: Northern Ter- of time constraints. ritory, East and West Alligator (QMB) AMNH American Museum of Natural History, To the 61 characters in Huber (2000), the New York following four characters were added: AMS Australian Museum, Sydney Character 62. ``Worm-shaped'' projection CLD Collection C. L. Deeleman-Reinhold, on genital bulb: (0) absent; (1) present. This re- Ossendrecht, Netherlands fers to a characteristic, more or less cylindrical, MNHN MuseÂum National d'Histoire Naturelle, membranous projection distally on the bulb in Paris most Wugigarra species (unshafted arrows in ®gs. QMB Queensland Museum, Brisbane 24, 64, 112, 157). This structure is either a syn- SAM South Australia Museum, Adelaide apomorphy of the genus with one or more rever- USNM National Museum of Natural History, sals, or it is the synapomorphy of only the core Washington group of Wugigarra (groups 1±3 under Speci®c WAM Western Australian Museum, Perth Relationships, p. 12). ZMH Zoologisches Museum, Hamburg Character 63. Weak zone on male palpal The methods used are described in Huber cymbium: (0) absent; (1) present. This refers to (2000). Eye measurements are Ϯ 5 ␮m. All a light (probably weakly sclerotized) zone dorsal- photos were done on a Hitachi S-4700 cold- ly on the male palpal cymbium (asterisks in ®gs. 201, 248, 265). This character is the proposed emission SEM. Localities are ®rst ordered by synapomorphy of Trichocyclus, and it is present territories and then roughly by their prox- in all species seen, except in one species assigned imity to the type locality. Cladogram analysis tentatively (T. watta). It is also present in Aucana was done with Clados, version 1.2 (Nixon, kaala Huber, a ninetine from New Caledonia, and 1992), and Winclada, version 0.9.9 ϩ (Nix- possibly also in Holocneminus. on, 1999). Character 64. ``Spermophora ¯ap'': (0) ab- sent; (1) present. This refers to a short ¯ap orig- RELATIONSHIPS inating at about the middle of the procursus ven- trally (``f'' in ®gs. 396, 410). This ¯ap is here The possible phylogenetic relationships of interpreted as not homologous to the ventral Australian genera were studied by simply hinged process of Micromerys and Metagonia, be- adding several Australian species to the ma- cause it is much shorter, apparently not hinged, trix in Huber (2000), and then submitting it and it is positioned more vertically relative to the to cladistic analysis. To the 61 taxa in Huber procursus (rather than parallel). As the name im- (2000), the following ten taxa were added: plies, this character seems to be restricted to Sper- mophora or a subgroup of Spermophora. It occurs 62. Trichocyclus arabana, n. sp.: Western both in the type species and in the two species Australia, Canning Stock Rte, Well 25 newly described. It occurs also in several undes- (WAM) cribed species from Kalimantan (Borneo), Sula- 63. Trichocyclus nigropunctatus Simon: wesi, Banda Islands (Moluccas), Sumatra, Phil- , Woodleigh Station ippines, and Sri Lanka (all in collection CLD). (WAM) Character 65. Posterior pocket or pockets on 64. Wugigarra kaurna, n. sp.: South Austra- female opisthosoma: (0) absent; (1) present. This character is here interpreted as synapomorphy of lia, Bunyeroo Gorge (SAM) Spermophora. Both newly described species have 65. Wugigarra sphaeroides (Koch): Queens- a median pocket between the epigynum and the land, Homevale (QMB) spinnerets (``p'' in ®gs. 399, 412; arrow in ®g. 66. Wugigarra bujundji, n. sp.: Queensland, 401). The type species, S. senoculata, has paired Home Rule (QMB) pockets that are situated on both sides of the spin- 2001 HUBER: PHOLCIDS OF AUSTRALIA 5 nerets. In this species, the long, hooked apophyses gain, of ALS piriform gland spigot loss ver- on the male genital bulbs are inserted into these sus gain, and of pseudosegmentation loss pockets during copulation (personal obs.). Pre- versus gain) left eight cladograms that dif- sumably, the hooked apophyses in other species fered only with respect to the New World (e.g., ``h'' in ®g. 396) serve the same function, clade, and only with respect to non-Austra- but only in S. senoculata are the apophyses so long that the pocket has to be split up and posi- lian taxa. One of these cladograms is shown tioned on both sides of the spinnerets. A median in appendix 2. However, no single cladogram pocket occurs also in several undescribed species satis®ed all constraining criteria above. For from Bali, Lombok and Sumbawa (Sunda Is- example, in all cladograms the AME were lands), Java, Philippines, and Papua New Guinea lost in the Pholcus group but were regained (all in collection CLD). in Pholcus and its closest relatives (clade 71 Instead of reproducing the entire matrix in appendix 2). Moreover, in the cladogram from Huber (2000), I give only the coding shown, the epiandrous spigots are lost in of characters for the new taxa (appendix 1). clade 135 (i.e., in the ancestor of holocne- The coding of the four new characters for the mines [clade 100] ϩ New World clade [clade ``old'' taxa follows logically from what is 134]), but are regained in Artema ϩ the Hol- said above: character 62, state (0) for all ocnemus group. Independent gains of these taxa; character 63, state (0) for all taxa; char- characters are possible theoretically, but a acters 64 and 65, state (0) for all taxa except slightly longer cladogram with only indepen- Spermophora senoculata. Apart from that, I dent losses would probably come closer to a have changed only one coding: character 17 phylogenetic tree than does the most parsi- (knob-shaped apophysis on epigynum) in monious cladogram. Spermophora senoculata was coded as pres- The groups consistently found in all to- ent in Huber (2000), but is now coded as pologies were (1) the Pholcus group sensu absent. Spermophora senoculata has either Huber, 1995 (clade 78), including most of the an extremely poorly developed, almost invis- genera present in Australia (Pholcus, Pan- ible knob or no knob at all. Only the study jange, Spermophora, Micromerys, and Beli- of closely related congeners might bring a sana); (2) Metagonia, an exclusively New satisfying solution to this point. World genus (clade 81); (3) ninetines (clade 90), a group with worldwide distribution, but without Australian representatives known; RESULTS AND DISCUSSION and (4) the New World clade (clade 134), The basic topologies obtained by NONA including the Australian genus Wugigarra (version 1.8; Goloboff, 1993) and Hennig86 (see discussion below). (version 1.5; Farris, 1988) with equally The only other Australian genus, Tricho- weighted characters were very similar to cyclus, is included in holocnemines (clade those obtained in the previous analysis with- 100: Holocnemus group sensu Timm, 1976 out Australian taxa (Huber, 2000). This sim- ϩ Artema, Physocyclus, Priscula, and Tri- ilarity included both the major clades, as well chocyclus) in the cladogram in appendix 2. as the ambiguity with respect to their inter- However, holocnemines were not consistent- relationships. As the worldwide cladogram ly resolved as a monophyletic group, but in seems thus not essentially improved, and the some topologies they were paraphyletic, with focus of this paper is rather the Australian the Holocnemus group branching off ®rst, fauna and its relationships, I only deal brie¯y then Artema, then Trichocyclus ϩ Physocy- with the overall topologies and concentrate clus, and ®nally Priscula as sister group of more on the positions of the Australian gen- the New World clade. These topologies were era. neither obviously better nor worse with re- Using NONA with hold/50, mult*100, spect to the constraints mentioned above. and amb- resulted in 81 most parsimonious Using Hennig86 with mh* and bb* re- cladograms of length 180 (CI ϭ 38; RI ϭ sulted in many more most parsimonious 77). Using the same criteria for tree selection cladograms with the same statistics, over- as in Huber (2000) (preference of AME loss ¯owing the tree buffer at 895 cladograms. versus gain, of epiandrous spigot loss versus The groups consistently found were the same 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 as above, with the same ambiguity with re- the present context. The only notable and spect to holocnemines and the same relation- possibly signi®cant difference from the ships of Australian genera. NONA cladogram in appendix 2 is that Me- In addition to the analyses using equally tagonia is nested within a paraphyletic Phol- weighted characters, two types of weighting cus group (clade 82), as sister taxon of Mi- were employed: implied weighting using cromerys, supported by the hinged process Pee-Wee (version 2.8; Goloboff, 1997), and on the procursus and the narrow sternum successive weighting using Hennig86 and (clade 78). The position of Trichocyclus as NONA. Successive weighting resulted in 69 sister group of ninetines is dubious and (Hennig86) and nine (NONA) most parsi- weakly supported; that is, by the narrow monious cladograms, but these were too ob- opening of the capsulate tarsal organ (coding viously much farther away from the true phy- ambiguous) and the presence of stridulation logeny to merit much attention. For example, on the male chelicerae (present also in sev- they required four to ®ve (Hennig86) or two eral holocnemines). Also, the sister group re- (NONA) independent gains of AME, and lationship of this clade with holocnemines, they nested ninetines within the New World resulting in clade 100, is not convincing. It clade, requiring the independent gain of ALS is based on the pointed cheliceral lamina, a piriform gland spigots and epiandrous spig- character that is dif®cult to code and that ots in ninetines. However, some of the within ninetines is restricted to Ibotyporan- groups found using equally weighted char- ga. acters were also found using successive The results con®rm the inclusion of most weighting; that is, the Pholcus group, Meta- Australian genera in the Pholcus group (as gonia, the Holocnemus group, and ninetines. proposed in Huber, 1995), but unfortunately Holocnemines were always paraphyletic, as they fall short of providing a convincing hy- was the New World clade. pothesis about the relationships of the two Implied weighting with Pee-Wee resulted dominant genera, Trichocyclus and Wugigar- in similarly unacceptable cladograms at most ra. Only the following can be said with some settings of the concavity (conc) constant. For con®dence: Trichocyclus is probably most example, at conc ϭ 1, 2, and 6, the absence closely related to several genera that together of epiandrous spigots and AME was primi- have a worldwide distribution, including tive for pholcids. Instead, the presence of Physocyclus in Central and North America, both characters is presumably the primitive Smeringopus in Africa, and Holocnemus, condition in haplogynes. At conc ϭ 3, 5, and Hoplopholcus, Crossopriza, and Artema in 6, ninetines were nested within the New northern Africa and the Middle East, and World clade, requiring the independent gain possibly also Priscula in South America. of ALS piriform gland spigots and epian- These genera share a brush of hairlike struc- drous spigots in ninetines. However, using tures distally on the procursus (not clearly conc ϭ 4 with hold/50 and mult*100 resulted present in the Holocnemus group), a tenden- in 43 most parsimonious cladograms of cy to reduce the pseudosegmentation of the length 184 (CI ϭ 37; RI ϭ 76), which failed tarsi (not in Trichocyclus), and narrowly only at one of the constraining criteria above: pointed cheliceral laminae. It is notable that AME were regained in Pholcus and its clos- these genera share some morphological and est relatives, as in the analyses using equally ecological characters that were not included weighted characters above. The only major in the matrix: all include mostly large phol- variation among these 43 cladograms con- cids (some Artema, Physocyclus, and Pris- cerned the basal polytomy of clade 131 in cula species are probably the largest phol- appendix 3, including Wugigarra and the cids), and most have the unusual tendency to (paraphyletic) New World clade. It was either be widespread in dry rather than in tropical a tetratomy (as shown in appendix 3) or a humid life zones (this is particularly true for trichotomy, with any of the three minor Trichocyclus, Physocyclus, Holocnemus, Ho- clades being the sister group of the large plopholcus, Crossopriza, and Artema). clade (clade 114). All other variation was More puzzling is the position of Wugigar- within this latter clade and is not relevant in ra. The inclusion within, or proximity to, the 2001 HUBER: PHOLCIDS OF AUSTRALIA 7

New World clade rests on a few characters, anthropic species seem to occur over the none of them convincing; that is, the absence whole continent. Some may actually be more of ALS piriform gland spigots (these have common in the tropical and subtropical areas been reduced at least ®ve times indepen- in the north, but I have not studied the dis- dently within pholcids; see Huber, 2000) and tributions of synanthropic species and will the exposed tarsal organ (the rim of the cup- not refer to them further. (2) Wugigarra (map shaped tarsal organ has been reduced at least 1) is extremely species rich in the east, rang- three times independently; see Huber, 2000). ing from Cape York Peninsula in northern Moreover, Wugigarra does not share with Queensland to Eyre Peninsula in South Aus- other New World clade genera the distinct tralia. The apparent distributional gap in Vic- retrolateral apophysis on the male palpal toria and southern New South Wales is prob- coxa, and it would be the only representative ably arti®cial, resulting in part from biases in in the New World clade to have cheliceral the collections studied and in part from the stridulation. Note that in the analyses using fact that these are among the areas with the weighted characters, Wugigarra was not con- highest habitat disturbance in Australia (Nix, sistently included in the New World clade 1981). Note, however, the absence of the ge- but was either sister group to the New World nus in the Northern Territory (with the ex- clade (which included ninetines, as men- ception of one doubtful record). (3) Tricho- tioned above) or had its origin in the same cyclus (map 2) covers the entire west of the polytomy as New World taxa. Thus, although continent, with gaps only in the driest deserts the exact relationships de®nitely remain ob- (Tanami Desert, Great Sandy Desert, Great scure, there seems to be some af®nity with Victoria Desert). The apparent gap in Arn- New World genera, and it is not without rea- hem Land is certainly arti®cial, while the gap son that previous authors included represen- in the far southwest may be real; that is, ex- tatives of Wugigarra in the New World ge- tensive collections in this region have pro- nus Psilochorus. duced no native pholcids (Mark Harvey, per- Apart from this apparent relationship with sonal commun.). Most interesting is the near New World genera in general, Wugigarra has absence of the genus in the east. Trichocy- a pair of characters that seem to place it close clus pustulatus in the Cairns area is a notable to three genera of holocnemines (Physocy- clus, Trichocyclus, and Artema): the dorsal exception, and more eastern localities will apophysis on the procursus, coupled with a probably be added in the future, but the con- ventral pouch on the same structure. Physo- trast with the distribution of Wugigarra is cyclus globosus is the only species of all striking. (4) The genera of the Pholcus group these genera in which the very peculiar func- (Pholcus, Spermophora, Panjange, Microm- tion of these characters has been established erys, Belisana) all have a northern and north- (Huber and Eberhard, 1997): they lock the eastern distribution, as is exempli®ed in map procursi together during copulation, ®tting 3 for Micromerys. Most of them do not reach the apophysis of one procursus into the farther south than the Cairns area, and Pan- pouch of the other one. This results in asym- jange seems to be restricted to the Cape York metrical insertion of the procursi, while all Peninsula. other pholcids studied have symmetrical in- What are the reasons for these distinct dis- sertion. The respective structures in Wugi- tributions? Both historical and contemporary garra, Trichocyclus, and Artema are suspi- factors are probably involved, and teasing ciously similar, and my prediction is that apart their respective importance would de- Wugigarra will eventually fall closer to these pend (1) on a more detailed knowledge of genera than to the New World clade. Note, the southeast Asian fauna from which several however, that the af®nity of Wugigarra to of the Australian genera seem to originate, New World taxa might remain intact, as Phy- and (2) on a more or less stable hypothesis socyclus is a New World genus. for the phylogenetic relationships of the taxa involved. The following can be said with BIOGEOGRAPHY some con®dence. First, the genera of the Australian pholcid genera and species Pholcus group all seem to originate from the show one of four distributions: (1) Most syn- north, having entered either via New Guinea 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Map 1. Distribution of the genus Wugigarra Map 2. Distribution of the genus Trichocylus. in Australia. This map includes both described This map includes both the described species and undescribed species, as well as unidenti®ed treated herein, as well as unidenti®ed female spec- female specimens in the collections studied. t ϭ imens in the collections studied. assigned tentatively; ? ϭ doubtful locality.

lems occur with respect to the origin and age or the Sunda Islands. This occurred probably of Micromerys. Micromerys is apparently a not earlier than the Pleistocene when sea lev- purely Australian genus, but the cladistic els were repeatedly signi®cantly lower. At analysis was not clearcut with respect to its this time, Australia was broadly connected to sister taxon. While it is most probably part New Guinea, and the Sunda and Sahul of the Pholcus group in the original sense shelves greatly facilitated dispersal over Ma- (i.e., excluding Metagonia), the possibility lesia (Malaya, Borneo, Sumatra, Java) that it is the sister taxon of the New World (Keast, 1981b). This northern origin is most genus Metagonia cannot be de®nitely dis- certainly true for Panjange, a genus that is missed. If this turns out to be the case, then so far only known from the Philippines, Bor- Micromerys would also be a relict Mesozoic neo, Celebes, and New Guinea (Deeleman- Reinhold, 1986a: map 5). It is also probably true for Belisana and Spermophora, both of which are exclusively Old World genera with wide distributions (not counting the synan- thropic S. senoculata), and in Australia these genera are restricted to the tropical north. Not so clear is the situation with Pholcus. Phol- cus is primarily an Old World genus, but the presence of various (undescribed) Pholcus species in caves of the eastern United States (R. Baptista, personal commun.) points to the possibility that Pholcus might be a pangaean genus dating back to at least the Jurassic, and Australian representatives might rather be pre-Eocene relicts than more recent northern intrusions. The scant data, however, suggest Map 3. Distribution of the genus Micromerys that Australian Pholcus species (at least P. in Australia. This map includes both the described tagoman) are most closely related to south- species treated herein, as well as unidenti®ed fe- east Asian species, which would rather hint male and juvenile specimens in the collections to a post-Miocene intrusion. Similar prob- studied. ? ϭ doubtful locality. 2001 HUBER: PHOLCIDS OF AUSTRALIA 9 element in Australia, rather than a Pleisto- cids have two origins: they are either relicts cene new tropical element. of Gondwanaland, with their presence in The two large Australian genera (Tricho- Australia dating back to the Mesozoic (Tri- cyclus, Wugigarra), on the other hand, are chocyclus, Wugigarra), or they are new trop- probably old (pre-Eocene) elements. Their ical elements having entered Australia from closest relatives, even though not de®nitely the north, probably not earlier than the Pleis- resolved (see Relationships above), seem to tocene. be genera from the New World (Physocyclus, Priscula), Africa (Smeringopus), and the KEY TO THE PHOLCIDS OF Middle Eastern±North African±Mediterra- AUSTRALIA nean region (Crossopriza, Artema, Holocne- This key is primarily designed to work mus, Hoplopholcus). Holocneminus, a south- well if males are available, but in most cases east Asian genus, probably belongs in this a female is suf®cient. In several cases the key group too. The striking east±west separation goes down to species level. Because all de- of Trichocyclus and Wugigarra (maps 1, 2) scribed autochthonous Australian species are may in part re¯ect ecological differences, but treated in this paper, I found it preferable in historical factors seem to play the major role. the other cases to end this key at the genus If Wugigarra were restricted geographically level and to refer for further determination to just for ecological and contemporaneous cli- the descriptive section. At this level, com- matic reasons, then it might be expected to paring illustrations and distributions is prob- occur at least in caves of the west and in ably a quicker and more reliable way to iden- typical refuge areas like tropical Kimberley tify species. and the Northern Territory, as well as in the far southwest (e.g., Stirling Ranges). This is 1. Six eyes; AME missing or reduced to pig- apparently not the case, however, and the real ment specks ...... 2 reason might rather be the marine subdivi- ± Eight eyes; AME always fully developed sion of the continent in the Cretaceous. At (with lenses) ...... 7 that time, extensive inroads of the sea in the 2(1). Opisthosoma roughly globular or higher than long ...... 3 area of what is now the Great Artesian Basin ± Opisthosoma at least three times longer separated the continent into a western and an than high ...... 6 eastern part (Brown et al., 1968), with a con- 3(2). Eyes on high eye turret; male clypeus nection remaining only in the south (which with frontal cuticular lobe; introduced was still connected to Antarctica). This sug- ...... culicinus gests a Cretaceous origin or ®rst major ra- ± Eyes not elevated or only slightly so 4 diation, with subsequent speciation facilitat- 4(3). Distance between posterior median eyes ed by the climatic ¯uctuations in the Pleis- (PME) more than three times their di- tocene. During this period, vegetation belts ameter; northeastern Queensland . . . repeatedly expanded and contracted, sug- ...... Spermophora ± Distance between PME less than two gesting repeated creation of isolates along times diameter of PME ...... 5 the Eastern Highlands after their creation in 5(4). Male chelicerae with one pair of pointed the Miocene (Keast, 1981a). distal apophyses and proximolateral A similar argument seems to apply to Tri- apophyses (®g. 423); male palpal tro- chocyclus, with the exception that the com- chanter with long apophysis (®g. 419); paratively more uniform western part of Aus- epigynum with scape (®g. 424) .... tralia offered less opportunities for speciation ...... Belisana australis and differentiation. This would explain the ± Male chelicerae with several apophyses comparative uniformity of the genus with re- on each side, without proximolateral spect to morphology. With the exception of apophyses (®g. 288); male palpal tro- chanter without apophysis (®g. 292); the species in the tropical North (Kimberley, epigynum without scape (®g. 289) . . . northern Northern Territory), most represen- ...... Trichocyclus watta tatives are quite similar. 6(2). Male genital bulb with only one projec- In sum, the present state of knowledge tion (e.g., ®gs. 302, 316); opisthosoma suggests that autochthonous Australian phol- worm-shaped (length Ͼ10ϫ diameter); 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

male chelicerae with only one pair of frontally; introduced ...... rounded light apophyses laterally (®g...... Artema atlanta 299); epigynum without scape ..... ± Conical projections on male chelicerae ...... Micromerys are apophyses; male chelicerae with ± Male genital bulb with two very long pro- stridulatory ridges laterally ..... 12 jections (``a'' and ``e'' in ®g. 385); op- 12(11). Male palpal femur with ventral apoph- isthosoma length ϳ5ϫ diameter medi- ysis (arrows in ®gs. 160, 167, 171); ally; opisthosoma drawn out into point females unknown; only in Cairns area dorsoposteriorly (®g. 378); male chelic- .. Wugigarra (idi, burgul, wanjuru) erae with only one pair of pointed light ± Male palpal femur without ventral apophyses laterally (®g. 380); epigyn- apophysis ...... 13 um with long scape (®g. 382) ..... 13(12). Male chelicerae with only two to three ...... Panjange mirabilis pairs of apophyses frontally (®gs. ± Male genital bulb with three projections 175, 185); male palpal femur only (``u'', ``e'', and ``a'' in ®g. 372); opis- slightly widened distally (®gs. 176, thosoma length ϳ3±4ϫ diameter me- 182) . . . Wugigarra (nauo, kalamai) dially, not drawn out into point dorso- ± Male chelicerae with four or more pairs posteriorly (®g. 363); male chelicerae of apophyses frontally; male palpal with one pair of rounded light apophy- femur in¯ated; female carapace with ses laterally and dark frontal apophyses median cone posteriorly; epigynum medially (®g. 369) ...... with forked apophysis frontally; intro- ...... Pholcus tagoman duced ...... Physocyclus globosus 7(1). Opisthosoma roughly globular or higher 14(7). Male chelicerae with proximolateral than long ...... 8 light apophyses (Pholcus) ...... 15 ± Opisthosoma at least three times longer ± Male chelicerae without proximolateral than high ...... 14 apophyses ...... 16 8(7). Procursus with long dorsal hinged pro- 15(14). Bulbal uncus wide, palpal trochanter cess; male chelicerae with proximola- apophysis short; epigynum with dis- teral light apophyses; pale, small phol- tinct ``knob'' frontally; introduced . . cid (ϳ1±1.5 mm total length); intro- ...... Pholcus phalangioides duced ...... Micropholcus fauroti ± Bulbal uncus narrow (®gs. 342, 358); ± Procursus without dorsal hinged process; male chelicerae without proximolateral palpal trochanter apophysis very long light apophyses; with dark markings, (®g. 343); epigynum with tiny or hid- rarely under 2 mm total length .... 9 den knob posteriorly (arrows in ®gs. 9(8). Genital bulb with worm-shaped projection 347, 348, 357, 361, 362); only ventrodistally (unshafted arrows in ®gs. Queensland ...... Pholcus (part) 24, 64, 106); epigynum often with me- 16(14). Male chelicerae with two pairs of dian pocket (e.g., ®gs. 26, 46, 66, 89) apophyses frontally; opisthosoma ...... Wugigarra (part) pointed posteriodorsally; introduced ± Genital bulb without worm-shaped projec- ...... Crossopriza lyoni tion ventrodistally; epigynum never ± Male chelicerae with one pair of apoph- with median pocket ...... 10 yses frontally; opisthosoma rounded 10(9). Male palpal cymbium with weak (light) posteriodorsally ...... 17 area dorsally (asterisks in ®gs. 201, 17(16). Male femur 1 with spines ventrally; 248, 265); male chelicerae without or male and female chelicerae with strid- with only one pair of cone-shaped pro- ulatory ridges; female palp enlarged; jections frontally; epigynum usually introduced .... Holocnemus pluchei with rounded median elevation frontal- ± Male femur 1 without spines ventrally; ly...... Trichocyclus (part) male and female chelicerae without ± Male palpal cymbium without weak stridulatory ridges; female palp not (light) area dorsally; male chelicerae enlarged ...... 18 with two or more pairs of cone-shaped 18(17). Genital bulb with two apophyses (®gs. projections frontally ...... 11 434, 435); epigynum as in ®g. 436; 11(10). Conical projections on male chelicerae introduced . . . Smeringopus pallidus are modi®ed hairs; male chelicerae ± Genital bulb with three apophyses (®gs. without stridulatory ridges laterally; 432, 433); epigynum as in ®g. 437; epigynum with a pair of dark humps introduced Smeringopus natalensis 2001 HUBER: PHOLCIDS OF AUSTRALIA 11

TAXONOMY line (e.g., ®gs. 5, 21, 33, 51, 79), sometimes WUGIGARRA, NEW GENUS with large humps (®gs. 34, 60, 124), modi- ®ed hairs (®g. 101), small cones (®gs. 110, TYPE SPECIES: Wugigarra tjapukai, new 134), or almost unmodi®ed (®gs. 115, 143). species. Male palps moderately to extremely large in ETYMOLOGY: The genus name is composed relation to overall size (e.g., ®gs. 1, 68, 158); of two words in Yidini, the aboriginal lan- coxa without retrolateral apophysis, trochan- guage of the Cairns-Yarrabah region: wugi, ter without apophyses, femur slightly to con- to shake, and garra, . It refers to the spicuously enlarged (e.g., ®g. 111), with re- shaking or whirling movements many phol- trolateral hump proximally, usually without cids (including representatives of this genus) ventral apophyses (W. bujundji, wunderlichi, make when disturbed. Gender feminine. and wulpura with large medioventral apoph- DIAGNOSIS: Small- to medium-sized (total ysis: ®g. 131; W. idi, burgul, and wanjuru length usually ϳ2±5 mm), pholcids with with small distal apophysis: ®gs. 160, 167, globular or higher-than-long opisthosoma, 171); trochanter triangular in lateral view; mostly with small AME (AME diameter usu- tibia simple, with 2 trichobothria in very ally 50±80% of PME diameter, in some spe- proximal position (e.g., ®gs. 35, 61, 111, cies up to 90%; in kalamai, 110%), appar- 171); cymbium with relatively simple pro- ently restricted to Australia or the Australian cursus that is usually provided with dorsal region. Distinguished from Trichocyclus apophysis and ventral pocket (``a'' and ``p'' (which is the only similar genus in Australia) in ®gs. 7, 61, 71, 103; apophysis always dis- by the absence of a weak zone dorsally on tinct, pocket often hardly recognizable), dis- the male cymbium; by the presence of a tally with membranous fringes, sometimes characteristic worm-shaped process on the also with hairlike structures (e.g., ®g. 47); bulb (unshafted arrows in ®gs. 24, 64, 96; bulb consisting of proximal globular part and missing only in some species that are as- distal sclerotized elements among which signed tentatively to this genus, see below); sperm duct opens without embolus (shafted by the presence of curved hairs on the legs; arrows in ®gs. 15, 24, 88, 96), usually with by the presence of only two spigots on the worm-shaped projection on ventral side (un- ALS (ϳ8±9 in Trichocyclus); by the pres- shafted arrows in ®gs. 24, 64, 96, 145, 157), ence of stridulatory ®les in females; and by missing only in two groups of closely related the epigynum that is often provided with a species (W. idi, burgul, wanjuru, and W. median pocket (e.g., ®gs. 26, 46, 76). nauo, kalamai). Legs usually long (leg 1 DESCRIPTION: Total length in males usually about 7±15 ϫ body length), medium-thin (tibia ϳ2±4 mm; only W. kaurna up to 5 mm, in 1 l/d ϳ55±90), leg 1 always the longest, legs W. burgul and wanjuru only ϳ1.5 mm. Car- 2 and 4 about same length, leg 3 shortest; apace oval, wider than long, with distinct with or without dark rings subdistally on fe- thoracic groove, often with median and lat- mur, patella ϩ tibia proximally, and tibia eral dark bands (e.g., ®gs. 3, 70), only in W. subdistally; tips of femora and tibiae often kaurna with three pairs of lateral spots loose- whitish; tarsal organ exposed (examined: W. ly connected to median spot by radial marks kaurna and sphaeroides: ®gs. 30, 49; see also (®g. 18), as in most Trichocyclus species. ®g. 84 in Huber, 2000). Legs usually without Eight eyes in conservative pattern, on mod- spines (present in W. undanbi and jiman), erately elevated ocular area. AME diameter with few vertical hairs, with curved hairs on ϳ30±90% of PME diameter (in W. kalamai tibiae and metatarsi, sometimes also on fem- 110%). Distance PME-ALE usually ϳ40± ora (curved hairs missing only in three spe- 80% of PME diameter, in W. arcoona only cies assigned tentatively: W. idi, burgul, and 25%, in W. eberhardi 100%. Clypeus usually wanjuru); retrolateral trichobothrium of tibia unmodi®ed (slightly modi®ed in W. tjapukai 1 usually at 5±15%, in W. jiman at 21%, in and mamu: ®gs. 2±4; and in W. bulburin: ®g. W. arcoona, nauo, and kalamai at 26±28%; 91). Male chelicerae always with stridulatory tarsus 1 with up to 45 pseudosegments, but ridges, otherwise extremely variable: often often only ϳ10±20 distal pseudosegments with pair of pointed apophyses near median easily visible in dissecting microscope. Op- 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 isthosoma either globular or higher than the procursus (dorsal apophysis and ventral long, gray with black spots and sometimes pouch or pocket) that is present also in some white spots dorsally; genital plate usually holocnemines (Physocyclus, Artema, Tricho- light brown, about rectangular, usually with cyclus). Cladistic analysis suggested that this distinct plate in front of spinnerets. Male similarity results from convergence. gonopore without epiandrous spigots (ex- Finally, it has been suggested (Deeleman- amined: W. sphaeroides, kaurna, and bu- Reinhold, 1995) that Australian ``Psilocho- jundji: ®gs. 32, 43, 137); ALS with only two rus'' (i.e., Wugigarra) are closely related to spigots (examined: W. sphaeroides, eberhar- Holocneminus. This is probably correct, but di, yawai, undanbi, bujundji, and wunderli- I have not seen enough material of Holoc- chi: ®gs. 28, 29, 48, 141); other spinnerets neminus to study it in detail. A preliminary typical for family. inclusion of an unidenti®ed Holocneminus Sexual dimorphism slight, females with species in the matrix resulted in paraphyletic shorter legs, often with larger (higher) opis- holocnemines, with Holocneminus as sister thosoma and darker sternum, chelicerae, and taxon of Wugigarra ϩ New World clade. palps; stridulatory ridges on chelicerae and SPECIFIC RELATIONSHIPS: Several opera- curved hairs on legs also in females. Only tional (not necessarily monophyletic) species W. nauo with enlarged female palps (®g. groups can be identi®ed by morphological 179). In some species female opisthosoma characters: (1) The most diverse and widely provided with median brown spot dorsofron- distributed tjapukai group is characterized in tally; corresponding side of thoracic groove the male by a pair of pointed apophyses close either unmodi®ed or slightly less deep. Epi- to the median line of the chelicerae (e.g., ®gs. gynum shape very variable, often with me- 5, 51, 79), and in the female by a median dian pocket (e.g., ®gs. 26, 57, 76) sometimes pocket on the epigynum (e.g., ®gs. 26, 46, situated on a long membranous scape (®gs. 76). This group includes the type species W. 41, 46); internally usually simple, with pair tjapukai and W. mamu, kaurna, sphaeroides, of relatively large pore plates (e.g., ®gs. 9, eberhardi, yawai, undanbi, jiman, and wiri. 42, 150); in W. bujundji and in wunderlichi Geographically, it covers most of the range complex, with median internal duct of un- of the genus, and might just be a paraphyletic known function (®gs. 149, 152). assemblage of primitive species. (2) A cer- MONOPHYLY: Most species included share tainly polyphyletic group of species with var- the worm-shaped projection ventrodistally on ious shapes of chelicerae and epigyna, but the bulb. Five species lacking this structure with bulbs that closely resemble those of the are assigned tentatively (W. idi, burgul, wan- ®rst group; this bulburin group includes W. juru, nauo, kalamai). bulburin, yirgay, arcoona, muluridji, and GENERIC RELATIONSHIPS: As discussed gia. (3) The probably monophyletic wunder- above (p. 6), the relationships of Wugigarra lichi group, with three described (W. bujund- are not clear. It might be either (1) part of ji, wunderlichi, wulpura) and at least one un- the New World group of genera, (2) the sister described species in northeastern Queens- taxon of this group, or (3) part of holocne- land, is characterized by the ventral hump on mines, a cosmopolitan group of genera. The the femur (®g. 131), the almost unmodi®ed cladograms in appendices 2 and 3 suggest the chelicerae (®gs. 134, 143), and the complex ®rst and second options, but this rests on structures in the female internal genitalia characters that show a relatively high degree (®gs. 149, 152). (4) A group lacking both the of homoplasy (reduction of ALS piriform worm-shaped projection ventrodistally on the gland spigots; reduction of the tarsal organ bulb and curved hairs on the legs, but as- rim, resulting in an exposed tarsal organ). signed to Wugigarra because of their overall Moreover, both the lack of a retrolateral similarity, includes three known species from apophysis on the male palpal coxa and the northeastern Queensland (W. idi, burgul, presence of cheliceral stridulation seem to ar- wanjuru). This idi group is probably mono- gue against these options. The third option phyletic (shape of ventral femur apophysis, was not supported by any cladistic analysis, shape of proximal palpal segments) and may but rests on the peculiar set of structures on be close to Holocneminus. (5) Two species 2001 HUBER: PHOLCIDS OF AUSTRALIA 13 from southern Australia that lack the worm- sented until 1980, ϳ15 were added by 1990, shaped process but have curved hairs (W. ϳ20 more by 2000. These numbers are ap- nauo, kalamai) are probably close relatives proximations because I did not make the ef- and share with W. arcoona (a more ``typical'' fort to sort the undescribed material into Wugigarra!) the distal position of the retro- morphospecies, but they strongly suggest lateral tibial trichobothrium and the geo- that the actual number of species may be graphic distribution (southern rather than well over 100. eastern Australia). NATURAL HISTORY: Most specimens were Wugigarra tjapukai, new species apparently collected in humid forests where Figures 1±12 they live mainly in the low vegetation and in Psilochorus sphaeroides (misidenti®cation; see the leaf litter. Only one species has been the Notes below): Jackson and Rowe, 1987; Jack- focus of detailed study: W. tjapukai (under son 1992; Jackson et al., 1992. Psilochorus sphaeroides: Jackson and Rowe, ?``Psilochorus'' sp. 1: Huber, 1998: ®g. 2M (see 1987; Jackson, 1992; Jackson et al., 1992). Notes below). This spider was observed to build webs sim- NOTES: Robert Jackson and coworkers did ilar to those of Pholcus phalangioides and to not publish detailed collection data of the invade contiguous webs, both alien and con- material they identi®ed as Psilochorus speci®c, presumably to steal prey. When at- sphaeroides (only ``Cairns'', or ``near tacked or disturbed, it started whirling, which Cairns''). A label accompanying the type reduced the ability of web-invading jumping material of the present species reads ``R. R. spiders to capture the pholcid. Whirling only Jackson, voucher specimens'', but these resulted from mechanical stimulation, not specimens were collected after the publica- from chemical stimuli from potential preda- tion of all the papers listed above. Neverthe- tors. less, since the only four vials in the QMB DISTRIBUTION: Presently known from Aus- containing specimens collected by R. Jack- tralia only. ``Psilochorus'' nigromaculatus son contain the present species, I ®nd it high- Kulczynski, 1911 from New Guinea might ly probable that this is actually the species be congeneric, but I have not been able to studied by Jackson and coworkers. see that species. The CLD collection has a Secondly, the species whose ``valve'' was male specimen of a probably undescribed studied in Huber (1998) is either conspeci®c species from Papua New Guinea (whose bulb or very close to the present species. (I copied seems to have a worm shaped process), and the erroneous label, saying ``Mittag Mittag'', three males of an undescribed species from which should be ``Millaa Millaa''.) Dumoga, Sulawesi, that seem very close to TYPE: Male holotype from Crystal Cas- the idi group above. Within Australia, the ge- cades near Cairns (16Њ58ЈS, 145Њ42ЈE), nus is remarkably restricted to the Great Di- Queensland, Australia; Dec. 1992 (R. R. viding Range in Queensland and New South Jackson), in QMB (S34679). Wales, with the exception of only four spe- ETYMOLOGY: Named for the Tjapukai, a cies west of longitude 140Њ in southern Aus- rainforest tribe in northeastern Queensland. tralia (map 1). The species name is a noun in apposition. COMPOSITION: The genus as construed here DIAGNOSIS: Distinguished from most con- includes a total of 22 described species, all geners by the modi®ed male clypeus (®gs. of which are treated below. Of these, two are 2±4), from all known congeners (including transferred from Psilochorus, the rest are the very similar Wugigarra mamu) by the new. The collections I have seen contain ϳ40 shapes of procursus tip (®gs. 10, 11) and dor- additional undescribed species, almost exclu- sodistal bulbal elements (®g. 12). sively members of the tjapukai group above. MALE (holotype): Total length 3.9, cara- Of this total of ϳ60 species, more than 50% pace width 1.84. Leg 1: 53.9 (13.1 ϩ 0.7 ϩ are known from the type locality only. More- 12.4 ϩ 19.7 ϩ 3.1), tibia 2: 8.1, tibia 3: 6.0, over, the number of new species per decade tibia 4: 8.4; tibia 1 l/d: 79. Habitus and pro- represented in the collections studied seems soma shape as in ®gs. 1±4. Carapace ochre to be increasing: to the ϳ25 species repre- with wide median and marginal brown 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 bands, thoracic furrow black (®g. 3). Ocular 145Њ54ЈE), 100 m elev., Queensland, Austra- area brown; distance PME-PME 0.135; di- lia; Oct. 17±24, 1981 (Earthwatch/QMB), in ameter PME 0.135; distance PME-ALE QMB (S27760). 0.105; diameter AME 0.105. Clypeus ochre ETYMOLOGY: Named for the Mamu, ab- yellow, distally modi®ed into black sclero- original rainforest dwellers in northeastern tized rim (®gs. 2±4); sternum dark brown, Queensland. The species name is a noun in with small yellowish speckles, margins ligh- apposition. ter (®g. 4). Chelicerae light brown, with pair DIAGNOSIS: Distinguished from most con- of black pointed apophyses medially and low geners by the modi®ed male clypeus (cf. ®gs. humps at their bases (®g. 5); apophyses not 2±4), from all known congeners (including visible in lateral view (®g. 1). Palps as in the very similar Wugigarra tjapukai) by the ®gs. 6±7, mostly ochre-yellow, only procur- shapes of procursus tip (®gs. 13, 14) and dor- sus and bulb partly brown to black; procur- sodistal bulbal elements (®gs. 15, 16). sus tip and bulb distinctive, as in ®gs. 6, 7, MALE (holotype): Total length 2.9, cara- 10±12. Legs ochre to light brown, without pace width 1.42. Leg 1: 38.9 (10.0 ϩ 0.5 ϩ dark rings, tips of femora and tibiae whitish; 9.6 ϩ 16.1 ϩ 2.7), tibia 2: 6.1, tibia 3: 4.4, curved hairs on tibiae and metatarsi; without tibia 4: 6.4; tibia 1 l/d: 85. Habitus and pro- spines and vertical hairs; retrolateral tricho- soma shape as in W. tjapukai (cf. ®gs. 1±4), bothrium of tibia 1 at 8%; tarsus 1 with Ͼ40 including color patterns and modi®ed clyp- distinct pseudosegments. Opisthosoma round- eus. Distance PME-PME 0.080; diameter ish, as in ®g. 1, gray, with black and white PME 0.135; distance PME-ALE 0.095; di- spots except ventrally; genital plate brown, ameter AME 0.105. Chelicerae and palps in about trapezoidal; brown plate in front of general identical to those in W. tjapukai (cf. spinnerets. ®gs. 5±7), but procursus tip and dorsodistal VARIATION: Tibia 1 in 7 males: 10.1±12.4 bulbal elements signi®cantly different (®gs. (xÅ ϭ 11.4). 13±16). Legs as in W. tjapukai, including FEMALE: In general very similar to male. curved hairs on tibiae and metatarsi; retro- Tibia 1 in 11 females: 8.8±10.9 (xÅ ϭ 9.7). lateral trichobothrium of tibia 1 at 7%; tarsus Epigynum as in ®g. 8, anterior plate laterally 1 with Ͼ40 distinct pseudosegments. Opis- dark brown, medially whitish, with transpar- thosoma identical to that in W. tjapukai (cf. ent scape, posterior plate light brown; dorsal ®g. 1). view as in ®g. 9. VARIATION: Tibia 1 in 7 males: 9.6±10.9 (xÅ DISTRIBUTION: Known from the Cairns area ϭ 10.4). (northeastern Queensland) and from one lo- FEMALE: In general very similar to male; cality about 300 km SSE of Cairns (map 4). tibia 1 in 8 females: 8.1±9.5 (xÅ ϭ 8.6). Epi- MATERIAL EXAMINED: AUSTRALIA: gynum externally not distinguishable from Queensland: Crystal Cascades near Cairns: that of W. tjapukai (cf. ®g. 8). Male holotype above, with 1( 3&, ``R. R. DISTRIBUTION: Known only from the Bel- Jackson, voucher specimens'' (QMB lenden Ker Range area near Cairns, north- S20897); same locality, Dec. 1982 (R. R. eastern Queensland (map 5). Jackson), 4( 4& (QMB S49973, 49963); MATERIAL EXAMINED: AUSTRALIA: Lamb Range (17Њ06ЈS, 145Њ37ЈE), Jan. 23, Queensland: Bellenden Ker Range, Cableway 1972 (N. Clyde Coleman), 1( (QMB Base Stn: Male holotype above, with 3& 1 S49979); Redlynch (16Њ53ЈS, 145Њ42ЈE), Jan. juvenile (QMB S34665); Bellenden Ker 1980 (R. R. Jackson), ``prey of Portia'', 5( Range, Westgid Creek (N Branch) (17Њ16ЈS, 5& (QMB S49812); Lower Mulgrave River 145Њ54ЈE), 100 m elev., Nov. 1, 1981 (Earth- (19Њ51ЈS, 147Њ10ЈE), Sept. 19, 1971 (N. watch/QMB), 1( (QMB S26257); Bellenden Clyde Coleman), 1( 1& (QMB S49883). Ker Range, 0.5 km S of Cable Tower No. 7 (17Њ16ЈS, 145Њ51ЈE), 500 m elev., Oct. 17± Wugigarra mamu, new species 24, 1981 (Earthwatch/QMB), 5( 3& (QMB Figures 13±16 S50181); same data but Oct. 25±31, 1981: TYPE: Male holotype from Bellenden 3& 5 juveniles (QMB S50182); same data Ker Range, Cableway Base Stn (17Њ16ЈS, but Nov. 1±7, 1981: 1( 2& (QMB S50239); 2001 HUBER: PHOLCIDS OF AUSTRALIA 15

Figs. 1±5. Wugigarra tjapukai, n. gen., n. sp., male. 1. Habitus, lateral view. 2±4. Prosoma, frontal, dorsal, and ventral views. 5. Chelicerae, frontal view. Scale lines: 1 mm (1), 0.5 mm (2±5).

Russell R. at Bellenden Ker Landing Wugigarra kaurna, new species (17Њ16ЈS, 145Њ57ЈE), 5 m elev., Nov. 1±9, Figures 17±32 1981 (Earthwatch/QMB), 1( 1& (QMB TYPE: Male holotype from Bunyeroo S50246); Graham Range (17Њ17ЈS, 145Њ58ЈE), Gorge (31Њ25ЈS, 138Њ34ЈE), ABC Range, 550 m elev., Nov. 1, 1995 (G. Monteith), Flinders Range National Park, South Austra- ``Pyrethrum, trees and logs'', 1( 2& (QMB lia, Australia; May 16, 1991 (D. Hirst), in S43364). SAM (N1999/717). 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 6±9. Wugigarra tjapukai, n. gen., n. sp. 6, 7. Left male palp, prolateral (6) and retrolateral (7) views; ``a'', ``p'' ϭ apophysis and pocket. 8, 9. Epigynum, ventral (8) and dorsal (9) views. Scale lines: 0.5 mm.

ETYMOLOGY: Named for the Kaurna, an DIAGNOSIS: Large species, distinguished aboriginal tribe from the Mount Lofty Range from congeners by the tip of the procursus area. The last woman survivor, Ivaritji, died (®gs. 22, 23), the distal bulbal elements (®gs. in 1931. The species name is a noun in ap- 24, 25), the shape of the epigynum (®g. 26), position. and the three pairs of lateral spots on the car- 2001 HUBER: PHOLCIDS OF AUSTRALIA 17

Figs. 10±16. Wugigarra tjapukai, n. gen., n. sp. (10±12), W. mamu, n. gen., n. sp. (13±16). 10, 11. Tip of left procursus, retrolateral (10) and prolateral (11) views. 12. Left genital bulb, dorsal (slightly retrolateral) view. 13, 14. Tip of left procursus, retrolateral (13) and prolateral (14) views. 15, 16. Left genital bulb, dorsal (15) and prolateral (16) views; shafted arrow: sperm duct opening; unshafted arrow: worm-shaped process. Scale lines: 0.2 mm. apace (®g. 18). The AMS has a possibly yellowish speckles. Chelicerae light ochre- closely related species from Capertee Valley brown, with pair of black pointed apophyses (ϳ33Њ10ЈS, 150Њ25ЈE), differing with respect medially and low humps at their bases (®g. to the procursus tip (AMS KS44134). 21); apophyses visible in lateral view (®g. MALE (holotype): Total length 4.8, cara- 17). Stridulatory ®les as in ®g. 31. Palps as pace width 2.03. Leg 1: 47.0 (12.8 ϩ 0.8 ϩ in ®g. 20, mostly ochre-yellow, only procur- 12.5 ϩ 18.1 ϩ 2.8), tibia 2: 8.9, tibia 3: 6.7, sus and bulb partly brown to black; procur- tibia 4: 8.9; tibia 1 l/d: 63. Habitus and pro- sus tip and bulb distinctive, as in ®gs. 22± soma shape as in ®gs. 17±19. Carapace pale 25. Palpal tarsal organ as in ®g. 30. Legs ochre with brown marks as in ®g. 18. Ocular ochre to light brown, dark rings on femora area pale ochre, slightly darker laterally; dis- subdistally, patellae ϩ tibiae proximally, and tance PME-PME 0.215; diameter PME tibiae subdistally; with many curved hairs on 0.145; distance PME-ALE 0.065; diameter tibiae and metatarsi 1±3; without spines and AME 0.095. Clypeus with pair of dark vertical hairs; retrolateral trichobothrium of stripes (®g. 19); sternum brown, with small tibia 1 at 8%. Tarsus 1 distally with ϳ15 fair- 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Australia: Flinders Range National Park: Male holotype above, with 5( 2& 2 juve- niles (SAM N1999/718±24); Flinders Range National Park, Heysen Range, Bunyeroo Gorge (31Њ25ЈS, 138Њ32ЈE), May 16, 1990 (D. Hirst), 3( 2& 1 juvenile in SAM (N1999/712±6); Gammon Ranges National Park, Arcoona Creek near Sambot WH (30Њ27ЈS, 139Њ02ЈE), at night on cliffs, May 4, 1989 (D. Hirst), 2( 2& (SAM N1999/ 706±9); rocky ridge NE of Arcoona Creek (30Њ26ЈS, 138Њ58ЈE), May 3, 1989 (D. Hirst), 1( 2 juveniles (SAM N1999/710); near Grindells Hut, Gammon Ranges National Park (30Њ29ЈS, 139Њ13ЈE), Apr. 1985 (B. Maps 4, 5. Distribution of Wugigarra species: Guerin), 1( (SAM N1999/711); Gammon tjapukai group. Ranges National Park, on cliff face, Wee- tootla Gorge (30Њ29ЈS, 139Њ14ЈE), Oct. 24, 1999 (D. Hirst), 1( 1 juvenile (SAM ly distinct pseudosegments; proximally pseu- NN9036); Mawson Plateau (30Њ07ЈS, dosegmentation dif®cult to see. Opisthosoma 139Њ25ЈE), Oct. 21±22, 1999 (D. Hirst), in as in ®g. 17, gray, with many black and some rocky creek bank, 1( 4& (SAM NN9031± white spots except ventrally. Genital plate 35); Alligator Gorge, Mt. Remarkable Na- only slightly darker, about trapezoidal; gon- tional Park (32Њ45ЈS, 138Њ03ЈE), on cliffs in opore without epiandrous spigots (®g. 32). gorge, Jan. 25, 1987 (D. Hirst), 1( 1& Plate in front of spinnerets indistinct. Two (SAM N1999/728±29); Melrose camping spigots on ALS (®g. 29, showing also the area (32Њ50ЈS, 138Њ11ЈE), Apr. 18, 1987 (D. usual two spigots on PMS). Hirst), 3& (SAM N1999/730±2); Spring Ck., VARIATION: There is an impressive north± near Melrose (32Њ50ЈS, 138Њ11ЈE), Apr. 21, south clinal variation, especially in sizeÐtib- 1973 (M. R. Gray), 1( (AMS KS51544); ia 1 in 15 males from north of latitude 32ЊS: Mambray Creek camp area (32Њ51ЈS, 10.5±13.7 (xÅ ϭ 11.3), in 9 males from south 138Њ00ЈE), from tree roots, steep creek bank, of 34ЊS: 7.3±8.9 (xÅ ϭ 8.3); the males from Oct. 4, 1988 (D. Hirst), 2( 1& (SAM between 32Њ and 34ЊS have intermediate val- N1999/725±7); Cave Cliff National Trust Re- ues: 8.9, 10.3, 10.3, 11.2. The same pattern serve, Hd Parcoola (33Њ37ЈS, 140Њ03ЈE), Apr. occurs in females (see below). In addition, 25, 1982 (J. J. Szent-Ivany), 1( 1& (SAM the number and size of teeth prolaterally on N1999/773±4); Cromer Conservation Park the male genital bulb follows a similar pat- (34Њ47ЈS, 138Њ59ЈE), Sept. 7, 1985 (B. Gue- tern: most specimens from northern popula- rin), under log, 1& (SAM N1999/733); Mt. tions have several teeth (as shown in ®gs. 24, Lofty Ranges, Castambul, Torrens Gorge 25); towards the south, these teeth get fewer (34Њ52ЈS, 138Њ46ЈE), Nov. 29, 1986 (D. and smaller, until they are completely absent. Hirst), 2( 2& (SAM N1999/740±3); Ade- FEMALE: In general very similar to maleÐ laide (34Њ56ЈS, 138Њ36ЈE), no date (R. H. tibia 1 in 8 females from north of latitude Pulleine), 1( 3& (poorly preserved) (SAM 32ЊS: 7.5±10.4 (xÅ ϭ 9.1), in 8 females from N1999/735±8); Adelaide, River Torrens south of 34ЊS: 5.5±8.0 (xÅ ϭ 6.7). Epigynum (34Њ56ЈS, 138Њ36ЈE), 1903 (R. H. Pulleine), as in ®g. 26, anterior plate laterally brown, 1& (SAM N1999/734); Adelaide, Windsor medially whitish with pocket on short trans- Gardens (34Њ56ЈS, 138Њ37ЈE), Dec. 1±16, parent scape; dorsal view as in ®g. 27. Spig- 1990 (D. Hirst), 1( (SAM N1999/739); Mt. ots on ALS as in male (®g. 28). Lofty Ranges, Belair (35Њ00ЈS, 138Њ38ЈE), DISTRIBUTION: Known from several local- Mar. 17, 1990 (L. N. Nicholson), 1( 1& ities in southeastern South Australia (map 4). (SAM N1999/748±9); same locality, Feb. 12, MATERIAL EXAMINED: AUSTRALIA: South 1990 (L. N. Nicholson), 1& (SAM N1999/ 2001 HUBER: PHOLCIDS OF AUSTRALIA 19

Figs. 17±21. Wugigarra kaurna, n. gen., n. sp., male. 17. Habitus, lateral view. 18, 19. Prosoma, dorsal and frontal view. 20. Left palp, retrolateral view. 21. Chelicerae, frontal view. Scale lines: 1 mm (18, 19), 0.3 mm (21).

750); same locality, Dec. 20, 1989 (L. N. 753); same locality, May 16, 1991 (L. N. Nicholson), 4& (SAM N1999/744±7); same Nicholson), in garden shed, 1& (SAM locality, Jan. 15, 1991 (L. N. Nicholson), 1& N1999/752); Mt Lofty Ranges, Loftia Park (SAM N1999/751); same locality, May 10, (35Њ02ЈS, 138Њ42ЈE), Sept. 14, 1989 (D. 1992 (L. N. Nicholson), 1& (SAM N1999/ Hirst), 1( (SAM N1999/755); same locality, 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 22±27. Wugigarra kaurna, n. gen., n. sp. 22, 23. Left procursus tip, retrolateral (22) and dorsal (23) views. 24, 25. Left genital bulb, prolateral (24) and dorsal (25) views; shafted arrow: sperm duct opening; unshafted arrow: worm-shaped process. 26, 27. Epigynum, ventral (26) and dorsal (27) views. Scale lines: 0.5 mm (24±27), 0.2 mm (22, 23).

Mar. 20, 1990 (D. Hirst), 4( 3& 2 juveniles TYPE: Koch (1867, 1872) described both (SAM N1999/756±63); Mt. Lofty Ranges, sexes from Rockhampton (23Њ22ЈS, 150Њ32ЈE), Hahndorf (35Њ02ЈS, 138Њ49ЈE), in fowl Queensland. This material seems to be lost, house, Dec. 1983 (B. Guerin), 3& (SAM but the ZMH has a male from Gayndah N1999/764±6); Mt. Lofty Ranges, Coroman- (ϳ270 km SE of Rockhampton) from the del Valley (35Њ02ЈS, 138Њ38ЈE), Apr. 1995 Museum Godeffroy collection that is very (L. N. Nicholson), 1& (SAM N1999/754); likely conspeci®c with Koch's original ma- Mt. Lofty Ranges near Clarendon (35Њ07ЈS, terial (see diagnosis below). 138Њ38ЈE), June 25, 1978 (A. F. Lees), 1( DIAGNOSIS: Closely related to W. yawai 2& (SAM N1999/768±70); Nappyalla and W. eberhardi, distinguished from both (35Њ20ЈS, 139Њ07ЈE), Mar. 1995 (J. Eckert), by the long transparent scape on the epigyn- 1& (SAM N1999/771); Snowgum Reserve, um (®gs. 41, 46) (Koch's [1872] ``weicher Carolyn Forest (37Њ56ЈS, 140Њ56ЈE), Apr. 20, . . . Fortsatz''), from the ®rst also by the 1979 (D. C. Lee), under Eucalyptus bark, 1& cheliceral apophyses not visible in lateral (SAM N1999/772). view (®g. 34) and the more slender procursus Wugigarra sphaeroides (Koch, 1867), (®g. 35), from the second also by the lateral new combination cones on the epigynum (®gs. 40, 41) (Koch's Figures 33±50 ``kegelfoÈrmige . . . HoÈckerchen''), and the Pholcus sphaeroides Koch, 1867: 193; 1872: ``stridulatory'' cone frontodorsally on the fe- 283±285, pl. 23, ®gs. 6, 6a±d. male opisthosoma (Koch's ``kleines HoÈck- Psilochorus sphaeroides: Simon, 1893: 481±482. erchen''). 2001 HUBER: PHOLCIDS OF AUSTRALIA 21

Figs. 28±32. Wugigarra kaurna, n. gen., n. sp. 28. Female ALS, with two spigots. 29. Male ALS (in front) and PMS (in back), with two spigots each. 30. Male palpal tarsal organ. 31. Stridulatory ®les on male chelicerae. 32. Male gonopore, without epiandrous spigots. Scale lines: 60 ␮m (31, 32), 20 ␮m (28, 29), 10 ␮m (30).

MALE (Homevale): Total length 3.8, cara- and procursus as in ®gs. 36±39. Brush of pace width 1.55. Leg 1: 36.1 (9.6 ϩ 0.7 ϩ hairlike structures on tip of procursus (®g. 9.6 ϩ 14.3 ϩ 1.9), tibia 2: 6.3, tibia 3: 4.5, 47). Legs ochre to light brown, with dark tibia 4: 6.4; tibia 1 l/d: 64. Habitus and pro- rings on femora subdistally, patellae ϩ tibiae soma shape similar to W. tjapukai (cf. ®gs. proximally, tibiae subdistally; tips of femora 1±4). Carapace ochre with wide brown me- and tibiae whitish; without spines and verti- dian band that is frontally widened, and with cal hairs; with curved hairs on tibiae and lateral bands. Ocular area brown; distance metatarsi 1±3 (few distally on femora and PME-PME 0.160; diameter PME 0.100; dis- proximally on metatarsi); retrolateral tricho- tance PME-ALE 0.060; diameter AME bothrium of tibia 1 at 10%; tarsus 1 distally 0.095. Clypeus brown, not modi®ed; sternum with ϳ16 distinct pseudosegments (®g. 44 brown, lateral margins ochre-yellow. Chelic- shows two from near the tip), proximally erae light brown with whitish humps and pseudosegmentation very indistinct. Opistho- black apophyses that are not visible in lateral soma shape as in W. tjapukai (cf. ®g. 1), view (®gs. 33, 34). Palps as in ®g. 35, bulb ochre gray with blackish spots dorsally. Gen- 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 ital plate brown; gonopore without epian- the type specimen and of many more phol- drous spigots (®g. 43). Brown plate in front cids from caves in New South Wales. of spinnerets; two spigots on ALS (®g. 48). DIAGNOSIS: Closely related to W. yawai VARIATION: Tibia 1 in 5 males: 8.1±10.0 (xÅ and sphaeroides, distinguished from both by ϭ 9.1). Measurements of Koch's ``type'' the low humps proximally on the male che- from Gayndah: total length 3.4, carapace licerae (®g. 52) and the absence of a spine width 1.6; leg 1: 10.0 ϩ 0.7 ϩ 9.7, metatar- on the genital bulb (®g. 56); from the ®rst sus and tarsus missing, tibia 2: 6.4, tibia 3: also by the cheliceral apophyses not visible 4.7, tibia 4: 6.7. in lateral view (®g. 52) and the more slender FEMALE: In general very similar to male, procursus (®g. 53), from the second by the but palps and chelicerae brown, and opistho- absence of a long transparent scape and lat- soma frontodorsally with unpaired sclero- eral cones on the epigynum (®g. 57). tized area, opposing part of thoracic furrow MALE (holotype): Total length 2.8, cara- that is less deep and more heavily sclerotized pace width 1.35. Leg 1: 26.1 (6.9 ϩ 0.5 ϩ than in male. Stridulatory ®les on chelicerae 7.1 ϩ 9.7 ϩ 1.9), tibia 2: 4.8, tibia 3: 3.5, as in ®g. 45. Palpal tarsal organ and palpal tibia 4: 4.8; tibia 1 l/d: 67. Habitus and pro- tarsus tip as in ®gs. 49 and 50. Epigynum as soma shape as in W. tjapukai (cf. ®gs. 1±4). in ®gs. 40 and 41, with anterior plate divided Carapace ochre with dark median line and into lateral brown parts with cones and me- spot behind ocular area, radial marks, and dian whitish part with long transparent scape light ochre spots laterofrontally. Ocular area provided with pocket (®gs. 41, 46); dorsal brown; distance PME-PME 0.175; diameter view as in ®g. 42. PME 0.065; distance PME-ALE 0.065; di- DISTRIBUTION: Known from several local- ameter AME 0.055. Clypeus brown, not ities between ϳ300 km N and 300 km S of modi®ed; sternum brown. Chelicerae brown Rockhampton, southeastern Queensland with low proximal humps and black apoph- (map 4). yses that are not visible in lateral view (®gs. MATERIAL EXAMINED: AUSTRALIA: 51, 52); with stridulatory ridges. Palps very Queensland: Rockhampton-Yeppoon (23Њ08ЈS, similar to W. sphaeroides (cf. ®g. 35), bulb 150Њ44ЈE), Oct. 29, 1973 (V. E. Davies), 1( and procursus as in ®gs. 53±56. Legs light 1& (QMB S50272); Rundle Range, ``site 5'' brown, without any rings; without spines and (23Њ39ЈS, 150Њ59ЈE), Mar. 24±31, 1975 (R. vertical hairs; with curved hairs on all tibiae Kohout, V. E. Davies), 1( 4& ϳ3 juveniles and metatarsi, and distally on femora; retro- (QMB S50162); Gayndah (25Њ37ЈS, lateral trichobothrium of tibia 1 at 7%; tarsus 151Њ37ЈE), ``No 11019'' (or ``1/019''?), no 1 distally with ϳ14 distinct pseudosegments, further data, 1( ``type'' in ZMH; Taroom, proximally pseudosegmentation very indis- Dawson R., Nathan Gorge (25Њ27ЈS, tinct. Opisthosoma shape as in W. tjapukai 150Њ08ЈE), Nov. 14, 1996 (P. Lawless), 1( (cf. ®g. 1), ochre with dark spots dorsally; 2& (QMB S37385, 37395); Homevale genital plate light brown, trapezoidal; brown (21Њ24ЈS, 148Њ33ЈE), Apr. 1±7, 1975 (R. Ko- plate in front of spinnerets. hout, V. E. Davies), 3( ϳ8& several juve- VARIATION: Tibia 1 in other males: 6.8, niles (QMB S50160); same locality, riverine 7.1, 8.7, 8.8. The male from Gecko Cave, rainforest, Apr. 1±7, 1975 (collector not giv- Gloucester, differs minimally with respect to en), 1& 1 juvenile (QMB S49976). the bulb (arrow in ®g. 53: this process is smaller; arrow in ®g. 56: more prominent Wugigarra eberhardi, new species hump). Figures 51±58 FEMALE: In general very similar to male. Epigynum without lateral elevations, with TYPE: Male holotype from Carrai Bat median pocket (®g. 57); dorsal view as in ®g. Cave, Stockyard Creek (30Њ59ЈS, 152Њ20ЈE), 58. New South Wales, Australia; Feb. 4, 1995 (S. DISTRIBUTION: Known from several local- Eberhard), in cave, ``space webs'', in AMS ities in eastern New South Wales (map 4). (KS65697). MATERIAL EXAMINED: AUSTRALIA: New ETYMOLOGY: Named for the collector of South Wales: Carrai Bat Cave, Stockyard 2001 HUBER: PHOLCIDS OF AUSTRALIA 23

Figs. 33±39. Wugigarra sphaeroides (Koch), n. comb., male. 33, 34. Chelicerae, frontal and lateral views. 35. Left palp, retrolateral view. 36, 37. Left procursus tip, retrolateral (36) and dorsal (37) views. 38, 39. Left genital bulb, dorsal (38) and prolateral (39) views. Scale lines: 0.3 mm (33±35, 38, 39), 0.2 mm (36, 37). 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 40±42. Wugigarra sphaeroides (Koch), n. comb., epigynum in ventral (40), lateral (41), and dorsal (42) views. Scale line: 0.5 mm (40±42).

Creek: Male holotype above, with 1( (AMS eral view (®g. 60) and the broad procursus KS49262); Youdale's Cave, Kunderang (®g. 61), from the ®rst also by the presence Brook (ϳ31Њ00ЈS, 152Њ12ЈE), Jan. 2, 1995 of a spine on the genital bulb (®g. 64); from (S. Eberhard), 1( 1& (AMS KS49264); Bat the second also by the absence of a long Cave, Yessabah (31Њ06ЈS, 152Њ41ЈE), Feb. transparent scape on the epigynum (®g. 66). 12, 1995 (S. Eberhard), dark zone, 2( 1 ju- MALE (holotype): Total length 2.4, cara- venile (AMS KS49259); N Plateau Rd ϳ3.5 pace width 1.23. Leg 1: 28.2 (7.5 ϩ 0.5 ϩ km from Plateau Beach Picnic Area 7.5 ϩ 10.8 ϩ 1.9), tibia 2: 4.8, tibia 3: 3.3, (31Њ11ЈS, 152Њ20ЈE), Mount Boss State For- tibia 4: 5.1; tibia 1 l/d: 75. Habitus and pro- est, Feb. 4±Apr. 9, 1993 (M. Gray, G. Cas- soma shape as in W. tjapukai (cf. ®gs. 1±4). sis), 1( (AMS KS42177); Gecko Cave, Carapace with dark pattern as in W. tjapukai Gloucester (ϳ32Њ02ЈS, 151Њ58ЈE), May 22, (cf. ®g. 3). Ocular area ochre; distance PME- 1995 (S. Eberhard), dark zone, 1( 1 juvenile PME 0.105; diameter PME 0.105; distance (AMS KS49266). PME-ALE 0.055; diameter AME 0.095. Clypeus with pair of dark stripes converging Wugigarra yawai, new species distally, not modi®ed; sternum dark brown Figures 59±67 except laterally. Chelicerae light brown with TYPE: Male holotype from Mt. Goonane- pair of high proximal humps and distinctive- man (25Њ26ЈS, 152Њ08ЈE), Queensland, Aus- ly projecting black apophyses that are visible tralia; Nov. 3±6, 1980 (R. Raven, V. E. Da- in lateral view (®gs. 59, 60), with stridula- vies), sieved litter, 670 m elev., in QMB tory ridges. Palps as in ®g. 61, bulb and pro- (S34675). cursus as in ®gs. 62±65. Legs light brown, ETYMOLOGY: Named for the Taribelang with slightly darker rings preceding whitish (also called Yawai), an aboriginal tribe in the tips of femora and tibiae; without spines and Bundaberg area. The species name is a noun vertical hairs; with curved hairs on all tibiae in apposition. and metatarsi 1±3; retrolateral trichoboth- DIAGNOSIS: Closely related to W. eberhardi rium of tibia 1 at 10%; tarsus 1 with Ͼ25 and sphaeroides, distinguished from both by pseudosegments, proximally pseudosegmen- the male cheliceral apophyses visible in lat- tation very indistinct. Opisthosoma shape as 2001 HUBER: PHOLCIDS OF AUSTRALIA 25

Figs. 43±50. Wugigarra sphaeroides (Koch), n. comb. 43. Male gonopore. 44. Male tarsus 1 near tip. 45. Stridulatory ridges on female right chelicera. 46. Scape on epigynum, ventrofrontal view. 47. Tip of procursus, showing brush of hairlike structures. 48. Male ALS, with two spigots. 49. Female palpal tarsal organ. 50. Tip of female palp. Scale lines: 50 ␮m (43±46), 20 ␮m (47, 48, 50), 5 ␮m (49). 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 51±58. Wugigarra eberhardi, n. gen., n. sp. 51, 52. Male chelicerae, frontal and lateral views. 53. Left procursus and bulb, retrolateral view. 54, 55. Tip of left procursus, retrolateral (54) and dorsal (55) views. 56. Left genital bulb, prolaterodorsal view. 57, 58. Epigynum, ventral (57) dorsal (58) views. Scale lines: 0.3 mm (51±53, 56±58), 0.2 mm (54, 55). Arrows point to variable structures. in W. tjapukai (cf. ®g. 1), gray with blackish of the bulb, distal to the large spine (arrow and white spots except ventrally; genital in ®g. 65), one or two additional tiny spines. plate brown; brown plate in front of spinner- FEMALE: In general very similar to male, ets. but chelicerae and palps dark brown, and op- VARIATION: Tibia 1 in 5 males: 6.0±7.5; isthosoma frontodorsally with unpaired scler- the male from Double Island is signi®cantly otized area, opposing part of thoracic furrow larger (tibia 1: 9.7), but is otherwise identi- that is less deep and more heavily sclerotized cal. Several males have on the prolateral side than in male. Tibia 1 in 8 females: 5.2±7.2; 2001 HUBER: PHOLCIDS OF AUSTRALIA 27

Figs. 59±65. Wugigarra yawai, n. gen., n. sp., male. 59, 60. Chelicerae, frontal and lateral views. 61. Left palp, retrolateral view; ``a'', ``p'' ϭ apophysis and pocket. 62, 63. Tip of left procursus, retrolateral (62) and dorsal (63) views. 64, 65. Left genital bulb, prolateral (64) and dorsal (65) views; unshafted arrow: worm-shaped process; shafted arrow: area where some males have 1±2 additional tiny spines. Scale lines: 0.4 mm (61, 64, 65), 0.2 mm (59, 60, 62, 63). 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 66±67. Wugigarra yawai, n. gen., n. sp. 66. Epigynum and spinnerets, ventral view; arrow: pocket. 67. Epigynum, dorsal view. Scale lines: 0.4 mm. female from Double Island signi®cantly larg- S50161); Dandabah, Bunya National Park er (tibia 1: 9.2). Epigynum with two pairs of (26Њ53ЈS, 151Њ37ЈE), Feb. 29, 1976 (un- lateral elevations and median pocket (®g. known collector), 1( 2& (QMB S50126); 66); in some females median light area less Marlaybrook, Bunya Mountains (26Њ54ЈS, diverging posteriorly; dorsal view as in ®g. 151Њ39ЈE), Mar. 6, 1976 (R. Raven, V. E. Da- 67. vies), 1( 1& (QMB S50141); Bunya Moun- DISTRIBUTION: Widely distributed in east- tain (26Њ54ЈS, 151Њ34ЈE), Sept. 4, 1974 (R. ern Queensland and northeastern New South Raven), 1( 1& (QMB S49816); Upper Wales, from ϳ20Њ±30ЊS (map 5). Brook®eld (27Њ50ЈS, 152Њ55ЈE), rainforest, MATERIAL EXAMINED: AUSTRALIA: April 29±May 13, 1981 (V. E. Davies, R. Queensland: Mt. Goonaneman: Male holotype Raven), 1( (QMB S49790); same locality above, with 1( 1& (QMB S50119); Good- and collectors, Mar. 18, 1981, 1& 1 juvenile night Scrub via Wallaville (25Њ05ЈS, (QMB S49965); Gold Creek Reservoir, 152Њ00ЈE), June 28, 1974 (J. Covacevich), Brook®eld (27Њ30ЈS, 152Њ55ЈE), Sept. 17, 1( (QMB S49886); Nipping Gully, ``site 2'' 1980 (R. Raven, V. E. Davies), 1& 2 juve- (25Њ40ЈS, 151Њ26ЈE), Aug. 21±Oct. 9, 1998 niles (QMB S 49977); Bahr's Scrub (G. B. Monteith), rainforest, 200 m elev., 1( (27Њ44ЈS, 153Њ10ЈE), May 23, 1981 (G. Mon- (QMB S49232); Double Island Pt., teith), 1( 1 juvenile (QMB S50194); Killar- ``near Little Freshwater Creek'' (25Њ56ЈS, ney, ``9 km out'' (ϳ28Њ20ЈS, 152Њ20ЈE), Aug. 153Њ11ЈE); Aug. 4, 1985 (J. Gallon, K. Sed- 10, 1997 (L. J. Boutin), under rocks and ler, R. Kropp), 1( 1& (QMB S49814); Coo- bark, 1( (QMB S40374); The Head, via Kil- loola (26Њ12ЈS, 153Њ03ЈE), Sept. 14, 1973 (R. larney (28Њ23ЈS, 152Њ19ЈE), Aug. 18±Nov. Raven), dead tree trunk, 1( 2 juveniles 17, 1974 (G. & S. Monteith), 1( (QMB (QMB S49885); Searys Scrub, Cooloola S50213); Kroombit Tops, Lower Dry Creek (26Њ12ЈS, 153Њ03ЈE), Feb. 6, 1976 (R. Raven, (24Њ24ЈS, 151Њ01ЈE), Dec. 9±19, 1983 (V. E. V. E. Davies), under logs, 4& (QMB Davies, J. Gallon), 1( 1& (QMB S49810); S50114); Mt. Coolum (26Њ34ЈS, 153Њ05ЈE), Kroombit Tops, 65 km SW of Gladstone Jan. 1984 (B. R. Jahake), 1( (QMB (24Њ22ЈS, 151Њ01ЈE), 1000±1100 m elev., 2001 HUBER: PHOLCIDS OF AUSTRALIA 29

Feb. 22±26, 1982, 1& (QMB S50314); Nob metatarsi 2 only, without vertical hairs; retro- Creek, By®eld (22Њ52ЈS, 150Њ37ЈE), Apr. 27, lateral trichobothrium of tibia 1 at 13%; tar- 1979 (G. B. Monteith), rainforest, sieved lit- sus 1 with Ͼ15 pseudosegments, proximally ter, 1& (QMB S50209); 5 km NW of Mt. pseudosegmentation very indistinct. Opistho- Macartney (20Њ49ЈS, 148Њ30ЈE), Apr. 21, soma shape as in ®g. 68, gray with blackish 1979 (G. B. Monteith), open forest, sieved spots except ventrally; genital plate large, litter, 480 m elev., 1( (QMB S50202); Pan- brown, trapezoidal; brown plate in front of danus Creek (20Њ48ЈS, 148Њ33ЈE), Cathu SF, spinnerets. Apr. 22, 1979 (G. B. Monteith), creek mar- VARIATION: Tibia 1 in 8 males: 3.6±4.8 (xÅ gin, 80 m elev., 1( 2& 2 juveniles (QMB ϭ 4.3). S50198, 50204, 50205). New South Wales: FEMALE: In general very similar to male, 0.5 km from Wheatly Creek Road on Camp but without spines on femora. Tibia 1 in 8 Creek Road (28Њ47ЈS, 152Њ19ЈE), Feb. 4± females: 2.4±3.6 (xÅ ϭ 2.8). Epigynum simple Apr. 9, 1993 (M. Gray, G. Cassis), 1( (AMS brown plate with posterior pocket (®g. 76); KS38374). dorsal view as in ®g. 77. DISTRIBUTION: Widely distributed in east- Wugigarra undanbi, new species ern Queensland, from ϳ20Њ±28ЊS (map 4). Figures 68±77 MATERIAL EXAMINED: AUSTRALIA: Queensland: Mt. Archer, Kilcoy: Male holo- TYPE: Male holotype from Mt. Archer, Kil- type above, with 1& in same vial; Mt. Mee, coy (26 57 S, 152 34 E), Queensland, Aus- Њ Ј Њ Ј ``site 2'' (27Њ05ЈS, 152Њ41ЈE), 550 m elev., tralia; Apr. 29, 1985 (J. Gallon), rainforest, June 26±Oct. 30, 1978 (G. B. Monteith), 1( 1500 ft. elev., in QMB (S34674). ϳ (QMB S50216); Neurum Creek, Mt. Mee ETYMOLOGY: Named for the Undanbi, an (27Њ05ЈS, 152Њ42ЈE), 210 m elev., Oct. 28, aboriginal tribe from northeastern Queens- 1977±Jan. 20, 1978 (G. B. Monteith), 1& land. The species name is a noun in apposi- (QMB S49975); Mt. Tenison Woods tion. (27Њ19ЈS, 152Њ45ЈE), rainforest, 750 m elev., DIAGNOSIS: Apparently closely related to W. eberhardi, yawai, and sphaeroides (note stick brushing, May 15, 1997 (G. B. Mon- the similarity in procursus and bulb shapes), teith), 1( 2& (QMB S43065); Mt. Glorious, distinguished by the smaller size (note the Hiller property (27Њ20ЈS, 152Њ46ЈE), Jan. 20± lack of overlap in tibia 1 length), the higher Apr. 26, 1978 (G. B. Monteith), 1& (QMB carapace and opisthosoma (®gs. 68, 69), and S49972); Mt. Glorious (27Њ20ЈS, 152Њ46ЈE), the simple ¯at epigynum with posterior Apr. 26, 1988 (P. Johns, V. E. Davies), 1( pocket (®g. 76). 1& (QMB S50302); same locality, Sept. 20, MALE (holotype): Total length 2.1, cara- 1979 (G. B. Monteith), rainforest, stick pace width 1.06. Leg 1: 16.6 (4.0 ϩ 0.4 ϩ brushing, 1( 1& 1 juvenile (QMB S50295); 4.4 ϩ 6.5 ϩ 1.3), tibia 2: 2.4, tibia 3: 1.9, Upper Brook®eld (27Њ30ЈS, 152Њ55ЈE), Sept. tibia 4: 2.5; tibia 1 l/d: 55. Habitus and pro- 7±Oct. 6, 1981 (V. E. Davies, R. Raven), soma shape as in ®gs. 68±70. Carapace rainforest, 1( (QMB S50294); same locality, ochre, with darker markings as in ®g. 70. Oc- Oct. 5, 1981 (V. E. Davies, R. Raven), rain- ular area ochre, laterally darker; distance forest, 2& 3 juveniles (QMB S50312); same PME-PME 0.135; diameter PME 0.095; dis- locality, Aug. 14, 1981 (R. Raven, V. E. Da- tance PME-ALE 0.045; diameter AME vies), litter, rainforest, 1( 2& 9 juveniles 0.055. Clypeus with wide dark mark; ster- (QMB S50301); same locality, Mar. 19, 1982 num brown. Chelicerae dark ochre, proxi- (R. Raven), litter, rainforest, 1& (QMB mally thicker but without hump, distally with S50309); same locality, Jan. 12, 1982 (R. Ra- pair of black apophyses not visible in lateral ven), litter, rainforest, 1& 1 juvenile (QMB view (®gs. 68, 69). Palps in general as in W. S50311); same locality, July 3, 1981 (V. E. yawai (cf. ®g. 61), bulb and procursus as in Davies, R. Raven, Ditter), rainforest, 2( 1& ®gs. 71±75. Legs light brown, without rings; (QMB S50313); 5 km NW of Mt. Macartney with single ventral row of spines on femora (20Њ49ЈS, 148Њ30ЈE), open forest, 480 m 1(ϳ25) and 2 (ϳ8), with curved hairs on elev., sieved litter, Apr. 21, 1979 (G. B. Mon- 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 68±75. Wugigarra undanbi, n. gen., n. sp., male. 68. Habitus, lateral view. 69, 70. Prosoma, frontal and dorsal views. 71. Left cymbium and procursus, retrolateral view; ``a'', ``p'' ϭ apophysis and pocket. 72, 73. Tip of left procursus, retrolateral (72) and dorsal (73) views. 74, 75. Genital bulb, dorsal (74) and prolateral (slightly dorsal) views. Scale lines: 1 mm (68), 0.5 mm (69, 70), 0.2 mm (71, 74, 75), 0.1 mm (72, 73).

teith), 4( (QMB S50191, 50199, 50214); Wugigarra jiman, new species Cape Hillsborough (20Њ55ЈS, 149Њ03ЈE), Figures 78±83 open forest, 10 m elev., sieved litter, Apr. 16, ?``Psilochorus'' sp. 2: Huber, 1998: ®g. 2L (see 1979 (G. B. Monteith), 1( (QMB S50215). Note below). 2001 HUBER: PHOLCIDS OF AUSTRALIA 31

Figs. 76±83. Wugigarra undanbi (76, 77), and W. jiman (78±83). 76. Epigynum and spinnerets, ventral view. 77. Epigynum, dorsal view. 78, 79. Male chelicerae, lateral and frontal views. 80, 81. Genital bulb, prolateral (80) and dorsal (slightly retrolateral) (81) views. 82. Left cymbium and procur- sus, retrolateral view. 83. Tip of left procursus, dorsal view. Scale lines: 0.2 mm.

?``Psilochorus'' sp.: Huber, 2000: ®gs. 84, 132, was collected from Magnetic Island, about 178 (see Note below). 700 km N of the localities listed below. TYPE: Male holotype from Taroom District NOTE: The species studied by Huber (25Њ25ЈS, 149Њ58ЈE), Queensland, Australia; (1998, 2000) is either conspeci®c or closely Nov. 12, 1996±Jan. 1997 (P. Lawless), vine related to the present species. However, it thicket on hill, in QMB (S37223). 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

ETYMOLOGY: Named for the Jiman, an ab- D. Cook), 1( (QMB S44351); same locality, original tribe from the Taroom area, Queens- Sept. 21±Dec. 19, 1997 (G. Monteith), 1( land. The species name is a noun in apposi- (QMB S44245). tion. DIAGNOSIS: Close relative of W. undanbi, Wugigarra wiri, new species easily distinguished from this and all other Figures 84±90 known congeners by the proximal position of the male cheliceral apophyses (®gs. 78, 79). TYPE: Male holotype from Finch Hatton MALE (holotype): Total length 1.9, cara- (21Њ09ЈS, 148Њ38ЈE), Queensland, Australia; pace width 0.97. Leg 1: 14.3 (3.7 ϩ 0.3 ϩ Apr. 7±14, 1975 (R. Kohout, V. E. Davies), 3.9 ϩ 5.3 ϩ 1.1), tibia 2: 2.3, tibia 3: 1.6, sheet web against rock, in QMB (S34678). tibia 4: 2.3; tibia 1 l/d: 59. Habitus and pro- ETYMOLOGY: Named for the Wiri, an ab- soma shape very similar to W. undanbi (cf. original tribe from middle-eastern Queens- ®gs. 68±70). Carapace ochre, with darker land. The species name is a noun in apposi- pattern similar to W. undanbi (cf. ®g. 70). tion. Ocular area ochre; distance PME-PME DIAGNOSIS: Distinguished from known 0.120; diameter PME 0.080; distance PME- congeners by the pair of apophyses distally ALE 0.040; diameter AME 0.055. Clypeus on the bulb (®gs. 87, 88) and by the shape with pair of brown marks; sternum brown. of the procursus tip (®gs. 84±86). Chelicerae brown, proximally with pair of MALE (holotype): Total length 3.1, cara- distinctive apophyses; with stridulatory ridg- pace width 1.48. Leg 1: 45.6 (11.7 ϩ 0.7 ϩ es (®gs. 78, 79). Palps in general similar to 11.7 ϩ 18.7 ϩ 2.8), tibia 2: 7.6, tibia 3: 5.5, W. yawai (cf. ®g. 61), bulb and procursus as tibia 4: 7.6; tibia 1 l/d: 88. Habitus and pro- in ®gs. 80±83. Legs light brown, tips of fem- soma shape as in W. tjapukai (cf. ®gs. 1±4). ora and tibiae whitish, preceded by darker rings; with single ventral row of spines on Carapace with dark pattern as in W. tjapukai femora 1 (ϳ20) and 2 (ϳ10), with curved (cf. ®g. 3). Ocular area brown; distance hairs on tibiae and metatarsi 1 and 2, without PME-PME 0.145; diameter PME 0.120; dis- vertical hairs; retrolateral trichobothrium of tance PME-ALE 0.095; diameter AME tibia 1 at 21%; tarsus 1 with Ͼ13 pseudos- 0.095. Clypeus with wide brown band, not egments, proximally pseudosegmentation modi®ed; sternum dark brown except later- very indistinct. Opisthosoma shape as in W. ally. Chelicerae brown, very similar to those undanbi (cf. ®g. 68), gray with blackish spots of W. tjapukai (cf. ®g. 5), with pair of black except ventrally; genital plate large, brown; apophyses whose tips are visible in lateral brown plate in front of spinnerets. view, without proximal humps, with stridu- VARIATION: Tibia 1 in other male: 3.5 latory ridges. Palps in general as in W. bul- (missing in others). burin (cf. ®g. 94), bulb and procursus as in FEMALE: Unknown (the single female sec- ®gs. 84±88. Legs light brown, tips of femora tioned in Huber [1998] had a pair of char- and tibiae whitish; without spines and verti- acteristic horns laterally on the epigynum). cal hairs; with curved hairs on tibiae and DISTRIBUTION: Known from two localities metatarsi 1 and 2; retrolateral trichobothrium in southeastern Queensland (map 5). The of tibia 1 at 9%; tarsus 1 with Ͼ25 pseudo- species might actually have a much wider segments, proximally pseudosegmentation distribution (see Note above). very indistinct. Opisthosoma shape as in W. MATERIAL EXAMINED: AUSTRALIA: tjapukai (cf. ®g. 1), ochre-gray with dark Queensland: Taroom District: Male holotype spots except ventrally; genital plate large, above; Mt. Rose via Taroom (25Њ25ЈS, brown; brown plate in front of spinnerets. 149Њ58ЈE), 260 m elev., vine scrub, Dec. 16, FEMALE: In general very similar to male, 1997±Mar. 4, 1998 (D. J. Cook), 1( (QMB but chelicerae and palps dark brown. Tibia S44355); Isla Gorge (25Њ16ЈS, 149Њ56ЈE), 8.4 1: 9.3 (missing in others). Epigynum brown, km SSW of lookout, vine scrub, 360 m elev., with posterior pocket (®g. 89); dorsal view Dec. 19, 1997±Mar. 3, 1998 (G. Monteith, as in ®g. 90. 2001 HUBER: PHOLCIDS OF AUSTRALIA 33

Figs. 84±90. Wugigarra wiri, n. gen., n. sp. 84. Left cymbium and procursus, retrolateral view. 85, 86. Tip of left procursus, prolateral (slightly dorsal) (85) and dorsal (86) views. 87, 88. Left genital bulb, retrolaterodorsal (87) and retrolateral (88) views; unshafted arrow: worm-shaped process; shafted arrow: sperm duct opening. 89, 90. Epigynum, ventral (89) and dorsal (90) views. Scale lines: 0.3 mm (84, 87±90), 0.2 mm (85, 86).

DISTRIBUTION: Known only from type lo- Wugigarra bulburin, new species cality in middle-eastern Queensland (map 5). Figures 91±100 MATERIAL EXAMINED: AUSTRALIA: Queensland: Finch Hatton: Male holo- TYPE: Male holotype from Bulburin type above, with 3& (QMB S50113). (24Њ30ЈS, 151Њ35ЈE), Queensland, Australia; 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Mar. 17±24, 1975 (R. Kohout, V. E. Davies), estry Nursery), NW of Bundaberg (24Њ31ЈS, in QMB (S34671). 151Њ29ЈE), rainforest, 580 m elev., Mar. 1975 ETYMOLOGY: Named for the type locality. (M. Gray, C. Horseman), 1( 1 juvenile The species name is a noun in apposition. (AMS KS6571); same data, 2& 1 juvenile DIAGNOSIS: Easily distinguished from all (AMS KS0099); same data, under rock and known congeners by the unique set of apoph- under logs, 3( (AMS KS6789). yses on the male chelicerae (®gs. 92, 93). MALE (holotype): Total length 1.9, cara- Wugigarra yirgay, new species pace width 1.00. Leg 1: 21.5 (5.3 ϩ 0.3 ϩ Figures 101±109 5.5 ϩ 8.8 ϩ 1.6), tibia 2: 3.1, tibia 3: 1.5, tibia 4: 3.0; tibia 1 l/d: 67. Habitus and pro- TYPE: Male holotype from Clifton Beach soma shape very similar to W. undanbi (cf. (16Њ46ЈS, 145Њ40ЈE), Queensland, Australia; ®gs. 68±70). Carapace ochre, with slightly 1971±1972 (N. Clyde Coleman), in QMB darker spot behind ocular area, small spot (S49884). posteriorly, and darker margins. Ocular area ETYMOLOGY: Named for the Irukandji (also ochre; distance PME-PME 0.120; diameter called Yirgay), an aboriginal tribe from the PME 0.095; distance PME-ALE 0.055; di- Cairns area, northeastern Queensland. The ameter AME 0.055. Clypeus with pair of species name is a noun in apposition. brown marks, distally with unsclerotized me- DIAGNOSIS: Easily distinguished from all dian projection (®g. 91); sternum light brown known congeners by the male chelicerae pro- laterally, darker longitudinal band medially. vided with several modi®ed hairs (®gs. 101, Chelicerae ochre, with set of very distinctive 102), and by the long thin procursus (®g. apophyses and stridulatory ridges (®gs. 92, 103). The QMB has a closely related unde- 93). Palps as in ®g. 94, bulb and procursus scribed species from Diamond Hill, Iron as in ®gs. 94±98. Legs ochre to light brown, Range (northern Queensland), with identical femora and tibiae with whitish tips preceded chelicerae, but with different procursus tip, by slightly darker rings; without spines and bulb, and epigynum (QMB S50263). vertical hairs; with curved hairs on tibiae and MALE (holotype): Total length 2.0, cara- metatarsi 1±3; retrolateral trichobothrium of pace width 0.97. Leg 1: 26.8 (6.9 ϩ 0.4 ϩ tibia 1 at 8%; tarsus 1 with Ͼ20 pseudo- 6.9 ϩ 10.7 ϩ 1.9), tibia 2: 4.0, tibia 3: 2.8, segments, proximally pseudosegmentation tibia 4: 4.1; tibia 1 l/d: 74. Habitus and pro- very indistinct. Opisthosoma shape similar to soma shape similar to W. tjapukai (cf. ®gs. W. undanbi (cf. ®g. 68), but not as high; gray 1±3). Carapace ochre, with slightly darker with large blackish spots except ventrally; median and lateral bands. Ocular area ochre; genital plate trapezoidal, brown; brown plate distance PME-PME 0.135; diameter PME in front of spinnerets. 0.085; distance PME-ALE 0.055; diameter VARIATION: Tibia 1 in 9 males: 5.5±5.7 (xÅ AME 0.055. Clypeus ochre, unmodi®ed; ϭ 5.6). sternum ochre-yellow. Chelicerae ochre, with FEMALE: In general similar to male, but dark brown modi®ed hairs, and stridulatory sternum without light lateral areas, and che- ridges (®g. 101). Palps as in ®g. 103, femur licerae and palps brown. Opisthosoma fron- without ventral apophysis, procursus tip and todorsally with unpaired sclerotized area, op- bulb as in ®gs. 103±107. Legs ochre-yellow, posing posterior part of thoracic furrow that with slightly darker rings on femora subdis- is less deep and more heavily sclerotized tally, patellae and tibiae proximally, tibiae than in male. Tibia 1 in 2 females: 4.1, 4.2. subdistally; tips of femora and tibiae whitish; Epigynum with posterior sclerotized scape, with curved hairs on metatarsi 1 and 2; with- apparently with pocket (®g. 99); dorsal view out spines and vertical hairs; retrolateral tri- as in ®g. 100. chobothrium on tibia 1 at 8%; tarsus 1 dis- DISTRIBUTION: Known only from Bulburin tally with ϳ15±20 fairly distinct pseudoseg- area, southeastern Queensland (map 6). ments, proximally pseudosegmentation MATERIAL EXAMINED: AUSTRALIA: dif®cult to see. Opisthosoma shape as in W. Queensland: Bulburin: Male holotype above, tjapukai (cf. ®g. 1), ochre gray with some with 5( 3& (QMB S49818); Bulburin (For- blackish spots except ventrally; genital plate 2001 HUBER: PHOLCIDS OF AUSTRALIA 35

Figs. 91±98. Wugigarra bulburin, n. gen., n. sp., male. 91. Prosoma, lateral view, showing slightly modi®ed clypeus. 92. Chelicerae, frontal view. 93. Cheliceral armature, oblique view. 94. Left palp, retrolateral view. 95, 96. Left genital bulb, dorsal (95) and prolateral (96) views; unshafted arrow: worm- shaped process; shafted arrow: sperm duct opening. 97, 98. Tip of left procursus, retrolateral (97) and dorsal (98) views. Scale lines: 0.5 mm (91, 94±96), 0.1 mm (92, 93, 97, 98). 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 99, 100. Wugigarra bulburin, n. gen., n. sp. Epigynum, ventral (99) and dorsal (100) views. Scale lines: 0.3 mm. hardly darker; light brown plate in front of above, with 1& in same vial; Endeavour spinnerets. Range 11 miles W of Cooktown (15Њ30ЈS, VARIATION: Tibia 1 in 2 other males: 8.1 145Њ06ЈE), Nov. 14, 1975 (R. Raven, V. E. (both). These two males are slightly larger Davies), litter, 1( (QMB S50269); Mt. Cook than the holotype, but appear identical in (15Њ30ЈS, 145Њ15ЈE), Nov. 14, 1975 (R. Ra- shape. ven, V. E. Davies), 1( 3 juveniles (QMB FEMALE: In general similar to male, but 50144). dark median band on carapace extends around ocular area, clypeus and sternum Wugigarra arcoona, new species darker (brown), and dark rings on legs more Figures 110±114 distinct. Epigynum very simple in ventral TYPE: Male holotype from Arcoona Bluff view (®g. 108), without pocket; dorsal view (30Њ26ЈS, 138Њ58ЈE), upper slopes at west as in ®g. 109. end of bluff, Gammon Ranges National Park, DISTRIBUTION: Known from three localities South Australia, Australia; May 3, 1989 (D. in northeastern Queensland (map 6). C. Lee), in SAM (N1999/781). MATERIAL EXAMINED: AUSTRALIA: ETYMOLOGY: Named for the type locality. Queensland: Clifton Beach: Male holotype The species name is a noun in apposition. DIAGNOSIS: Easily distinguished from all known congeners by the male chelicerae having several small cones proximally and a pair of larger apophyses distally (®g. 110), and by the shape of the procursus tip (®g. 114) and the bulb (®gs. 112, 113). MALE (holotype): Total length 2.3, cara- pace width 1.03. Leg 1: 17.1 (4.7 ϩ 0.4 ϩ 4.9 ϩ 5.9 ϩ 1.2), tibia 2: 3.4, tibia 3: 2.5, tibia 4: 3.3; tibia 1 l/d: 57. Habitus and pro- soma shape as in W. undanbi (cf. ®gs. 68± 70), but carapace less high. Carapace ochre- yellow, with slightly darker band around oc- ular area. Ocular area light ochre; distance PME-PME 0.115; diameter PME 0.085; dis- tance PME-ALE 0.020; diameter AME Maps 6, 7. Distribution of Wugigarra species: 0.080. Clypeus with slightly darker wide bulburin group (map 6, on left), and wunderlichi mark; sternum whitish. Chelicerae ochre to group (map 7, on right). light brown, proximally much wider than 2001 HUBER: PHOLCIDS OF AUSTRALIA 37

Figs. 101±109. Wugigarra yirgai, n. gen., n. sp. 101. Male chelicerae, frontal view. 102. Modi®ed hairs on male chelicerae. 103. Left male palp, retrolateral view; ``a'', ``p'' ϭ apophysis and pocket. 104, 105. Tip of left procursus, retrolateral (104) and prolateral (105) views. 106, 107. Left genital bulb, prolateral (106) and dorsal (107) views. 108, 109. Epigynum, ventral (108) and dorsal (109) views. Scale lines: 0.4 mm (103, 106±109), 0.2 mm (101, 104, 105), 0.05 mm (102). Unshafted arrows: worm- shaped process. 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 110±114. Wugigarra arcoona, n. gen., n. sp., male. 110. Chelicerae, frontal view. 111. Left palp, retrolateral view. 112, 113. Left genital bulb, prolateral (112) and dorsal (113) views. 114. Tip of left procursus, retrolateral view. Scale lines: 0.4 mm (111±113), 0.2 mm (110, 114). Unshafted arrows: worm-shaped process. distally, with some black cones, with stridu- dif®cult to see. Opisthosoma shape as in W. latory ridges, distally with pair of inward- undanbi (cf. ®g. 70), ochre gray with some facing apophyses (®g. 110). Palps as in ®g. blackish spots except ventrally; genital plate 111, femur without ventral apophysis, pro- and plate in front of spinnerets not darker. cursus tip and bulb as in ®gs. 112±114. Legs VARIATION: Tibia 1 in 4 other males: 3.1, ochre-yellow, darker rings hardly visible; 4.3, 4.7, 4.8. with curved hairs on tibia 1 and metatarsi 1 FEMALE: The SAM has a female specimen and 2; without spines and vertical hairs; ret- from ``base of Beda Hill, South Gap Station'' rolateral trichobothrium on tibia 1 at 28%; (31Њ51ЈS, 137Њ37ЈE) that might be conspecif- tarsus 1 distally with ϳ10 indistinct pseu- ic (SAM N1999/784). It has a very distinc- dosegments, proximally pseudosegmentation tive epigynum with a sclerotized scape. 2001 HUBER: PHOLCIDS OF AUSTRALIA 39

DISTRIBUTION: Known from two areas ble (femora subdistally, tibiae proximally and more than 2000 km apart: Gammon Ranges subdistally), tips of femora and tibiae whit- in South Australia, and Melville Island in ish; curved hairs on tibiae 1±3 and all meta- Northern Territory (map 6). I personally ®nd tarsi; without spines and vertical hairs; retro- the Melville record dubious, and suggest that lateral trichobothrium on tibia 1 at 7%; tarsus one question it until further material is found. 1 distally with ϳ25 fairly distinct pseudo- MATERIAL EXAMINED: AUSTRALIA: South segments, proximally pseudosegmentation Australia: Gammon Ranges, Arcoona Bluff: dif®cult to see. Opisthosoma shape as in W. Male holotype above; same data, 1( (SAM tjapukai (cf. ®g. 1), dark gray with blackish N1999/778); Arcoona Creek near Elephant spots except ventrally; genital plate and plate Hill (30Њ26ЈS, 138Њ59ЈE), May 5, 1989 (D. in front of spinnerets light brown. Hirst), 1( (SAM N1999/782); 3.3 km SSW VARIATION: Tibia 1 in other male: 9.3. of Welcome Well, Arcoona Station (31Њ17ЈS, FEMALE: In general very similar to male, 137Њ02ЈE), Nov. 10±14, 1996 (Stony Desert but chelicerae and palps brown. Tibia 1 in Survey), 1( (SAM N1999/783). Northern two females: 6.5, 7.8. Epigynum large, Territory: Melville Island, Pickertaramoor brown, without pocket (®g. 121); dorsal view (ϳ11Њ46ЈS, 130Њ53ЈE), Jan. 16, 1990 (M. J. as in ®g. 122. Tyler), 1( in SAM (N1999/860). DISTRIBUTION: Known only from type lo- cality in northeastern Queensland (map 6). Wugigarra muluridji, new species MATERIAL EXAMINED: AUSTRALIA: Figures 115±122 Queensland: Bakers Blue Mt.: Male holotype above, with 1( 2& 1 juvenile (QMB 47811). TYPE: Male holotype from Bakers Blue Mt., 17 km W of Mt. Molloy (16Њ42ЈS, 145Њ10ЈE), Queensland, Australia; Jan. 8±9, Wugigarra gia, new species 1990 (ANZSES), 800±1000 m elev., in Figures 123±130 QMB (S34680). TYPE: Male holotype from Mt. Dryander ETYMOLOGY: Named for the Muluridji, an aboriginal tribe from northeastern Queens- (20Њ15ЈS, 148Њ33ЈE), Queensland, Australia; land. The species name is a noun in apposi- Nov. 21, 1992 (G. Monteith, G. Thompson, tion. H. Janetzki), 700 m elev., in QMB (S49559). DIAGNOSIS: Easily distinguished from most ETYMOLOGY: Named for the Gia, an ab- known congeners by the male chelicerae that original tribe from the Proserpine area. The have no modi®cation other than a proximal species name is a noun in apposition. pair of humps (®g. 115); W. wunderlichi has DIAGNOSIS: Easily distinguished from all similar chelicerae but extremely different known congeners by the male chelicerae palps (e.g., femora with ventral apophyses). with their large, light, bi®d protrusions (®gs. MALE (holotype): Total length 2.8, cara- 123, 124) and by the tip of the procursus and pace width 1.35. Leg 1: ϳ36.5 (9.6 ϩ 0.4 ϩ the bulb (®gs. 125±127). 9.6 ϩϳ14.5 ϩϳ2.4), tibia 2: 6.3, tibia 3: MALE (holotype): Total length 3.7, cara- 4.5, tibia 4: 6.4. Habitus and prosoma shape pace width 1.74. Leg 1 missing, tibia 2: 7.9, as in W. tjapukai (cf. ®gs. 1±4). Carapace tibia 3: 5.7, tibia 4: 7.7. Habitus and prosoma ochre-brown, with darker median band and shape as in W. tjapukai (cf. ®gs. 1±4). Car- lateral margins. Ocular area brown; distance apace ochre, medially and laterally slightly PME-PME 0.135; diameter PME 0.105; dis- darker. Ocular area slightly darker than car- tance PME-ALE 0.065; diameter AME apace; distance PME-PME 0.185; diameter 0.065. Clypeus ochre-brown; sternum light PME 0.105; distance PME-ALE 0.055; di- brown. Chelicerae light brown, with proxi- ameter AME 0.065. Clypeus ochre, unmod- mal bulge and stridulatory ridges (®g. 115); i®ed; sternum ochre-yellow. Chelicerae hairs on bulge slightly stronger than others. ochre to light brown, with pair of light pro- Palps in general as in W. tjapukai (cf. ®gs. trusions and stridulatory ridges (®gs. 123, 6, 7), procursus and bulb as in ®gs. 116±120. 124). Palps in general similar to W. yawai Legs ochre-brown, darker rings hardly visi- (cf. ®g. 161), but retrolateral apophysis on 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 115±122. Wugigarra muluridji, n. gen., n. sp. 115. Male chelicerae, frontal view. 116. Left cymbium and procursus, retrolateral view. 117. Left procursus, dorsal view. 118±120. Left genital bulb, retrolateral (118), dorsal (119), and prolateral (120) views. 121, 122. Epigynum, ventral (121) and dorsal (122) views. Scale lines: 0.3 mm. femur without distal projection; procursus retrolateral trichobothrium on tibia 2 at 11%; and bulb as in ®gs. 125±127. Legs light tarsus 2 distally with ϳ14 quite distinct pseu- brown, without darker rings, tips of tibiae dosegments, proximally pseudosegmentation whitish; with curved hairs on tibiae and dif®cult to see. Opisthosoma shape as in W. metatarsi; without spines and vertical hairs; tjapukai (cf. ®g. 1), ochre-gray, covered with 2001 HUBER: PHOLCIDS OF AUSTRALIA 41

Figs. 123±130. Wugigarra gia, n. gen., n. sp. 123, 124. Male chelicerae, frontal and lateral views. 125. Procursus and genital bulb, retrolateral view. 126. Genital bulb, prolateral view. 127. Tip of pro- cursus, dorsal view. 128. Epigynum, ventral view. 129. Epigynum and spinnerets, lateral view. 130. Epigynum, dorsal view. Scale lines: 0.5 mm (128±130), 0.3 (123±127). blackish spots except ventrally; genital plate sofrontally with small transverse plate, but and plate in front of spinnerets light brown. opposing side of carapace apparently not FEMALE: In general very similar to male, modi®ed. Epigynum as in ®gs. 128, 129; dor- but sternum much darker; opisthosoma dor- sal view as in ®g. 130. 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

DISTRIBUTION: Known only from type lo- tally with ϳ18 quite distinct pseudosegments cality in middle-eastern Queensland (map 6). (®g. 136 shows two near the tip), proximally MATERIAL EXAMINED: AUSTRALIA: pseudosegmentation not visible in dissecting Queensland: Mt. Dryander: Male holotype microscope. Tarsal claws as in ®g. 142. Op- above, with 1& in same vial. isthosoma similar to W. tjapukai, slightly longer, ochre-gray, with blackish spots ex- Wugigarra bujundji, new species cept ventrally. Genital plate brown, about Figures 131±142, 148±150 trapezoidal; gonopore without epiandrous TYPE: Male holotype from Mt. Finnigan spigots (®g. 137). Brown plate in front of (15Њ49ЈS, 145Њ17ЈE), Queensland, Australia; spinnerets. Nov. 9, 1974 (L. R., V. E. Davies, D. Joffe), VARIATION: Tibia 1 in 9 males: 9.7±11.9 (xÅ on tree trunks, 3200±3600Ј elev., in QMB ϭ 10.5). Some males have an additional (S49974). small spine where the arrow points in ®g. ETYMOLOGY: Named for the Kokobujundji 135. (also called Bujundji), an aboriginal tribe FEMALE: In general very similar to male, from the Mt. Finnigan area. The species but chelicerae and palps darker brown; fewer name is a noun in apposition. stridulatory ridges than male (®g. 139). Tibia DIAGNOSIS: Closely related to W. wunder- 1 in 13 females: 7.3±10.0 (xÅ ϭ 8.7). Epigyn- lichi and wulpura, distinguished by the pair um as in ®g. 148, light to dark brown; dorsal of apophyses distally on the male chelicerae view and cleared ventral view as in ®gs. 149 (®g. 134), by the tip of the procursus (com- and 150. Two spigots on ALS (®g. 141). pare ®gs. 133, 147, 155), and by the bulbal DISTRIBUTION: Known from several local- apophyses (compare ®gs. 131 and 135 with ities in the Cairns area, northeastern Queens- 114±146). The QMB also has a very close land (map 7). undescribed relative from Spear Creek, MATERIAL EXAMINED: AUSTRALIA: northeastern Queensland (QMB 50139, Queensland: Mt. Finnigan: Male holotype 50117), which differs only in details of pro- above, with 1& in same vial; Mt. Finlay cursus and bulb shape. (15Њ49ЈS, 145Њ21ЈE), Nov. 29, 1975 (R. M., MALE (holotype): Total length 3.3, cara- V. E. Davies), 2( 4& 1 juvenile (QMB pace width 1.58. Leg 1: 39.5 (10.0 ϩ 0.7 ϩ S49978); Twelve Mile Scrub (15Њ50ЈS, 9.9 ϩ 16.1 ϩ 2.8), tibia 2 missing, tibia 3: 145Њ19ЈE), Nov. 22±27, 1975 (collector not 5.1, tibia 4 missing; tibia 1 l/d: 74. Habitus given), complex mesophyll vine forest on and prosoma shape as in W. tjapukai (cf. ®gs. granite, 1( 1& 1 juvenile (QMB S50140); 1±4). Carapace ochre, with wide median and Gap Creek (15Њ50ЈS, 145Њ19ЈE), Nov. 19, marginal brown bands. Ocular area brown; 1974 (D. Joffe), 1( 2& (QMB S50127); Mt. distance PME-PME 0.145; diameter PME Hartley (15Њ46ЈS, 145Њ20ЈE), 1600 ft elev., 0.145; distance PME-ALE 0.095; diameter Nov. 6, 1974 (J. C., D. Joffe, V. E. Davies), AME 0.105. Clypeus ochre, unmodi®ed; 1& (QMB S50138); Home Rule, Granites sternum brown, lateral margins lighter. Che- Track (15Њ44ЈS, 145Њ18ЈE), tangle web licerae light brown, with pair of small apoph- against tree trunks, 1200' elev., Nov. 16, yses distally and low humps frontally (®g. 1974 (D. Joffe, V. E. Davies), 5( 6& (QMB 134); stridulatory ridges as in ®g. 138. Strid- S50261); same locality, Nov. 11, 1974 (J.C., ulatory pick is a modi®ed hair proximally on D.J., K.Mc.D.), 3& (QMB S50121); Home palpal femur (®g. 140). Palps as in ®gs. 131 Rule (15Њ44ЈS, 145Њ18ЈE), in base of dead and 132, mostly ochre-yellow, only procur- palm frond, Nov. 13, 1974 (D. Joffe), 1( 1& sus and bulb partly brown to black; femur (QMB S50115); Mt. Hedley, The Hummock with distinct ventral apophysis (®g. 131); (15Њ44ЈS, 145Њ17ЈE), ®ne shawl web in hol- procursus tip and bulb distinctive, as in ®gs. low, Oct. 12, 1974 (V. E. Davies), 2( 2& 131Ð133 and 135. Legs ochre-brown, with- (QMB S50116); Mt Fisher, 7 km SW of Mil- out rings, tips of femora and tibiae whitish; laa Millaa (17Њ33ЈS, 145Њ34ЈE), Apr. 27±29, curved hairs on tibia 1 and all metatarsi; 1982 (G. Monteith, D. Yeates, D. Cook), without spines and vertical hairs; retrolateral 1050±1100 m elev., 1( 1& 2 juveniles trichobothrium of tibia 1 at 5%; tarsus 1 dis- (QMB S50238). 2001 HUBER: PHOLCIDS OF AUSTRALIA 43

Figs. 131±135. Wugigarra bujundji, n. gen., n. sp., male. 131, 132. Left palp, prolateral (131) and retrolateral (132) views. 133. Tip of left procursus, dorsal view. 134. Chelicerae, frontal view. 135. Left genital bulb, dorsal view; arrow: area where some males have a small additional spine. Scale lines: 0.5 mm (131, 132), 0.3 mm (133±135). 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 136±142. Wugigarra bujundji, n. gen., n. sp. 136. Pseudosegmentation of male tarsus 1. 137. Male gonopore. 138, 139. Stridulatory ridges on male (138) and female (139) chelicerae. 140. Stridu- latory pick on female palpal femur. 141. Female ALS with two spigots. 142. Tarsal claws on male tarsus 1. Scale lines: 60 ␮m (138, 139), 30 ␮m (136, 137, 142), 15 ␮m (141), 5 ␮m (140). 2001 HUBER: PHOLCIDS OF AUSTRALIA 45

Figs. 143±147. Wugigarra wunderlichi (Deeleman-Reinhold), n. comb., male. 143. Chelicerae, fron- tal view. 144. Left procursus and genital bulb, retrolateral view. 145, 147. Left genital bulb, prolateral (145) and dorsal (147) views; arrow: worm-shaped process. 146. Tip of left procursus, dorsal view. Scale lines: 0.3 mm (143±145, 147), 0.1 mm (146).

Wugigarra wunderlichi TYPES: Male holotype from Cape Tribula- (Deeleman-Reinhold, 1995), tion (16Њ05ЈS, 145Њ26ЈE), Queensland, Aus- new combination tralia; July±Aug. 1992 (J. Wunderlich), in Figures 143±147, 151±153 QMB (S51022), examined. One male and one female paratypes, same collection data, Psilochorus wunderlichi Deeleman-Reinhold, in QMB (male) and Collection Deeleman- 1995: 37±38, ®gs. 16±21. Reinhold (female), not examined. 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 148±153. Wugigarra bujundji (148±150) and W. wunderlichi (151±153), epigyna. 148, 151. Ventral views. 149, 152. Cleared ventral views. 150, 153. Dorsal views. Scale lines: 0.3 mm (148±153).

DIAGNOSIS: Closely related to W. bujundji and by the bulbal apophyses (compare ®gs. and wulpura, distinguished from the ®rst by 145 and 147 with 156 and 157). the absence of apophyses on the male che- MALE (type locality, QMB S50304): Total licerae (®g. 143), from the second by the tip length 2.8, carapace width 1.42. Leg 1: 40.4 of the procursus (compare ®gs. 144 and 155), (10.4 ϩ 0.5 ϩ 10.0 ϩ 16.8 ϩ 2.7), tibia 2: 2001 HUBER: PHOLCIDS OF AUSTRALIA 47

6.6, tibia 3: 4.7, tibia 4: 6.8; tibia 1 l/d: 75. niles (QMB S50304); Cape Tribulation, Pil- Habitus and prosoma shape as in W. tjapukai grim Sands (16Њ04ЈS, 145Њ28ЈE), Aug. 27, (cf. ®gs. 1±4). Carapace ochre, with wide 1988 (J.C., T.C., R. Raven), 1( 1& (QMB median and marginal brown bands. Ocular S13924). area brown; distance PME-PME 0.105; di- ameter PME 0.135; distance PME-ALE Wugigarra wulpura, new species 0.095; diameter AME 0.080. Clypeus brown, Figures 154±157 unmodi®ed; sternum dark brown. Chelicerae brown, unmodi®ed except pair of frontal TYPE: Male holotype from 4.5±5 km W of bulges, with stridulatory ridges (®g. 143). Cape Tribulation (16Њ05ЈS, 145Њ26ЈE), Top Palps in general as in W. bujundji (cf. ®gs. Camp, Queensland, Australia; 760±780 m 131, 132), femur apophysis slightly thinner elev., Oct. 1±6, 1982 (G. Monteith, D. Yea- and more proximal; procursus tip and bulb tes, G. Thompson), in QMB (S34662). distinctive, as in ®gs. 144±147. Legs light ETYMOLOGY: Named for the Wulpura, ab- brown, without darker rings, tips of femora original rainforest dwellers in northeastern and tibiae whitish; curved hairs on tibiae 1 Queensland. The species name is a noun in and 2 and on metatarsi 1, 2, and 4; without apposition. spines and vertical hairs; retrolateral tricho- DIAGNOSIS: Closely related to W. wunder- bothrium of tibia 1 at 7%; tarsus 1 distally lichi and bujundji, distinguished from both with ϳ25 quite distinct pseudosegments, by the two long dorsal projections on the proximally pseudosegmentation not visible procursus (®gs. 154, 155), and by the bulbal in dissecting microscope. Opisthosoma sim- apophyses (®gs. 156, 157). ilar to W. tjapukai, but slightly longer; ochre- MALE (holotype): Total length 3.1, cara- gray, with many blackish and some white pace width 1.61. Leg 1: 38.5 (9.5 ϩ 0.5 ϩ spots except ventrally; genital plate brown, 9.5 ϩ 15.9 ϩ 3.1), tibia 2: 6.7, tibia 3: 4.8, about trapezoidal; dark brown plate in front tibia 4: 7.0; tibia 1 l/d: 68. Habitus and pro- of spinnerets. soma shape as in W. tjapukai (cf. ®gs. 1±4). VARIATION: Tibia 1 in 3 other males: 9.2, Carapace ochre, with wide median and mar- 9.3, 9.6 (holotype); carapace width in holo- ginal brown bands. Ocular area brown; dis- type: 1.28. tance PME-PME 0.145; diameter PME FEMALE: In general very similar to male, 0.145; distance PME-ALE 0.120; diameter but chelicerae and palps darker brown; strid- AME 0.095. Clypeus ochre, unmodi®ed; ulatory ®les on chelicerae present, not absent sternum brown, lateral margins lighter. Che- as described by Deeleman-Reinhold (1995); licerae light brown, similar to those of W. tibia 1 in 2 females: 7.6, 8.9. Epigynum as wunderlichi (cf. ®g. 143), with low humps in ®gs. 151±153, light to dark brown; dorsal frontally and stridulatory ridges; without view and cleared ventral view as in ®gs. 152 apophyses. Palps in general as in W. bujundji and 153. (cf. ®gs. 131, 132), but femur more slender DISTRIBUTION: Known only from the Cape and ventral apophysis smaller; procursus tip Tribulation area, northeastern Queensland and bulb distinctive, as in ®gs. 154±157. (map 7). Legs light brown, without dark rings, tips of MATERIAL EXAMINED: AUSTRALIA: femora and tibiae whitish; with curved hairs Queensland: Cape Tribulation: Male holotype on tibiae 1 and 2 and metatarsi 1±3; without above; 5 km W of Cape Tribulation (16Њ05ЈS, spines and vertical hairs; retrolateral tricho- 145Њ26ЈE), 780 m elev., Sept. 28, 1982 (G. bothrium of tibia 1 at 6%; tarsus 1 distally Monteith, D. Yeates, G. Thompson), 1& 1 with ϳ25 quite distinct pseudosegments, juvenile, (QMB S50298); 3.0 km W of Cape proximally pseudosegmentation not visible Tribulation (16Њ05ЈS, 145Њ27ЈE), 500 m elev., in dissecting microscope. Opisthosoma sim- Oct. 2, 1982 (G. Monteith, D. Yeates, G. ilar to W. tjapukai (cf. ®g. 1), slightly longer, Thompson), 1( 1& (QMB S50318); 1.5 km ochre-gray, with blackish spots except ven- NW of Cape Tribulation (16Њ05ЈS, trally; genital plate brown, about trapezoidal; 145Њ28ЈE), sea level, Oct. 8, 1982 (G. Mon- brown plate in front of spinnerets. teith, D. Yeates, G. Thompson), 4( 2 juve- FEMALE: In general very similar to male, 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 154±157. Wugigarra wulpura, male. 154. Tip of left procursus, prolateral view. 155. Left cymbium and procursus, retrolateral view. 156, 157. Left genital bulb, dorsal (156) and retrolateral (157) views; arrow: worm-shaped process. Scale line: 0.5 mm (154±157).

but chelicerae and palps darker brown; tibia ETYMOLOGY: Named for the Idindji (also 1 in single known female: 8.9. Epigynum ex- called Idi), aboriginal rainforest dwellers ternally not distinguishable from that of W. from the Cairns area, northeastern Queens- wunderlichi (cf. ®g. 151). It is possible that land. The species name is a noun in apposi- this female is in fact W. wunderlichi, which tion. occurs in the same area. DIAGNOSIS: Easily distinguished from most DISTRIBUTION: Known only from type lo- congeners by the male chelicerae (several cality near Cape Tribulation, Queensland cones; ®g. 161); from W. burgul (which has (map 7). very similar chelicerae) by the procursus MATERIAL EXAMINED: AUSTRALIA: (longer and very different tip: ®gs. 160, 162), Queensland: 4.5±5 km W of Cape Tribulation: and especially by the bulb with its prominent Male holotype above, with 1( 1& 2 juve- prolateral armature of black apophyses (®gs. niles (QMB S50293). 159, 163). The QMB has a very close un- described relative from Mt. Bartle-Frere, 0.5 Wugigarra idi, new species km N of S Peak (17Њ24ЈS, 145Њ49ЈE) (QMB Figures 158±163 S49721), with different procursus tip and TYPE: Male holotype from Bellenden Ker bulbal apophyses. Range, Summit TV Station (17Њ16ЈS, MALE (holotype): Total length 2.0, cara- 145Њ37ЈE), Queensland, Australia; 1560 m pace width 1.10. Leg 1: 18.3 (4.7 ϩ 0.3 ϩ elev., dung trap in rainforest, Nov. 1±7, 1981 4.8 ϩ 6.9 ϩ 1.6), tibia 2: 2.9, tibia 3: 2.1, (Earthwatch/Queensland Museum), in QMB tibia 4: 3.0; tibia 1 l/d: 63. Habitus as in ®g. (S50235). 158. Prosoma shape similar to W. undanbi 2001 HUBER: PHOLCIDS OF AUSTRALIA 49

(cf. ®gs. 69, 70), but carapace less elevated; DIAGNOSIS: Easily distinguished from most carapace ochre, medially with slightly darker congeners by the male chelicerae (several broad band. Ocular area light brown; dis- cones; ®g. 165); from W. idi (which has very tance PME-PME 0.120; diameter PME similar chelicerae) by the procursus (shorter 0.085; distance PME-ALE 0.040; diameter and very different tip; ®gs. 167±169), and by AME 0.040. Clypeus with wide brown mark; the simpler bulb (compare ®gs. 159 and sternum dark brown. Chelicerae light brown, 166). with several small cones (®g. 161), proximal MALE (holotype): Total length 1.4 (1.6 cones on elevation (®g. 158). Palps as in ®gs. with clypeus), carapace width 0.72. Leg 1 159 and 160; femur with distinct retrolatero- missing, tibia 2: 2.24, tibia 3: 1.56, tibia 4: ventral apophysis (arrow in ®g. 160); bulb 2.24. Habitus as in ®g. 164. Prosoma shape and procursus tip as in ®gs. 159, 160, 162, similar to W. undanbi (cf. ®gs. 69, 70), but 163. Legs light brown, with indistinct darker carapace less elevated; carapace ochre, with- rings subdistally on femora and tibiae; tips out dark marks. Ocular area ochre; distance of femora and tibiae whitish; without spines, PME-PME 0.095; diameter PME 0.075; dis- curved and vertical hairs; retrolateral tricho- tance PME-ALE 0.035; diameter AME bothrium of tibia 1 at 14%; tarsus 1 with Ͼ15 0.025. Clypeus slightly darker than carapace; pseudosegments, proximally pseudosegmen- sternum brown. Chelicerae light brown, with tation very indistinct. Opisthosoma shape as several small cones (®g. 165), proximal in ®g. 158, gray with blackish spots except cones on slight elevation. Palps as in ®gs. ventrally; genital plate large, light brown, 166 and 167, femur with distinct retrolatero- trapezoidal; light brown plate in front of ventral apophysis (arrow in ®g. 167), pro- spinnerets. cursus tip as in ®gs. 168 and 169. Legs VARIATION: Tibia 1 in 2 other males: 5.2 ochre, with indistinct darker rings subdistally (both). on femora and tibiae; without spines, curved, FEMALE: Unknown. and vertical hairs (legs 1 missing). Opistho- DISTRIBUTION: Known only from Bellen- soma shape as in ®g. 164, gray with large den Ker Range, northeastern Queensland blackish spots except ventrally; genital plate (map 8). light brown, trapezoidal; light brown plate in MATERIAL EXAMINED: AUSTRALIA: front of spinnerets. Queensland: Bellenden Ker Range, Summit FEMALE: Unknown. TV Station: Male holotype above; same data, 1( (QMB S50237); same locality, Oct. 28, DISTRIBUTION: Known only from type lo- 1983 (G. Monteith, D. Yeates, G. Thomp- cality in northeastern Queensland (map 8). son), 1( (poorly preserved) (QMB S50253); MATERIAL EXAMINED: AUSTRALIA: Bellenden Ker Range, Cable Tower 3 Queensland: 3.0 km W of Cape Tribulation: (17Њ16ЈS, 145Њ51ЈE), 1054 m elev., Oct. 25± Male holotype above. 31, 1981 (Earthwatch/Queensland Museum), 1( (QMB S27997). Wugigarra wanjuru, new species Figures 170±172 Wugigarra burgul, new species TYPE: Male holotype from east margin of Figures 164±169 Palmerston National Park (17Њ37ЈS, 145Њ46ЈE), TYPE: Male holotype from 3.0 km W of Queensland, Australia; 400 m elev., Dec. 10, Cape Tribulation (16Њ05ЈS, 145Њ27ЈE), 1995±Feb. 7, 1996 (G. Monteith, D. Cook), Queensland, Australia; 500 m elev., Oct. 2, in QMB (S43258). 1982 (G. Monteith, D. Yeates, G. Thomp- ETYMOLOGY: Named for the Wanjuru, ab- son), in QMB (S34659). original rainforest dwellers in the Innisfail ETYMOLOGY: In Yidini, the aboriginal lan- and Babinda area. The species name is a guage of the Cairns-Yarrabah region, burgul noun in apposition. is a mythical being, believed to be able to DIAGNOSIS: Easily distinguished from con- take on any form. The species name is a geners by the male chelicerae provided with noun in apposition. a few small cones (®g. 170), and by the long 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 158±160. Wugigarra idi, male. 158. Habitus, lateral view. 159, 160. Left palp, prolateral (159) and retrolateral (160) views; arrow: femur apophysis. Scale lines: 1 mm (158), 0.5 mm (159, 160). dorsodistal projection on the procursus (®g. (cf. ®g. 164). Prosoma shape similar to W. 171). undanbi (cf. ®gs. 69, 70), but carapace less MALE (holotype): Total length 1.5, cara- elevated; carapace ochre, only medially and pace width 0.64. Leg 1: 13.97 (3.41 ϩ 0.27 around ocular area slightly darker. Ocular ϩ 3.61 ϩ 5.51 ϩ 1.17), tibia 2: 1.95, tibia 3: area ochre, slightly darker than carapace; dis- 1.32, tibia 4 missing. Habitus as in W. burgul tance PME-PME 0.095; diameter PME 2001 HUBER: PHOLCIDS OF AUSTRALIA 51

Figs. 161±163. Wugigarra idi, male. 161. Chelicerae, frontal view. 162. Tip of left procursus, re- trolateral view. 163. Left genital bulb, ``distal'' view; arrow: sperm duct opening. Scale lines: 0.3 mm.

0.065; distance PME-ALE 0.025; diameter without spines, curved, and vertical hairs AME 0.025. Clypeus slightly darker than (legs 4 missing). Opisthosoma shape as in W. carapace; sternum brown. Chelicerae light burgul (cf. ®g. 164), gray with blackish spots brown, with some small cones and stridula- except ventrally; genital plate light brown, tory ridges (®g. 170). Palps as in ®g. 171; trapezoidal; light brown plate in front of femur with distinct retrolateroventral apoph- spinnerets. ysis (arrow in ®g. 171); bulb as in ®gs. 171 FEMALE: Unknown. and 172. Legs pale ochre, with indistinct DISTRIBUTION: Known only from type lo- darker rings subdistally on femora and tibiae; cality in northeastern Queensland (map 8). MATERIAL EXAMINED: AUSTRALIA: Queensland: Palmerston National Park: Male holotype above.

Wugigarra nauo, new species Figures 173±181 TYPE: Male holotype from Kirton Point, Port Lincoln (34Њ43ЈS, 135Њ52ЈE), South Australia, Australia; Dec. 17, 1981 (D. Hirst), in SAM (N1999/855). ETYMOLOGY: Named for the Nauo, a now extinct aboriginal tribe that originally inhab- ited the coastal scrub gum tree forest country in South Australia. The species name is a noun in apposition. DIAGNOSIS: Distinguished from the closely Map 8. Distribution of Wugigarra species: related W. kalamai by the much larger size, species assigned tentatively (idi group and nauo by the apophyses on the male chelicerae group). (compare ®gs. 175 and 185), and by details 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 164±169. Wugigarra burgul, male. 164. Habitus, lateral view. 165. Chelicerae, frontal view. 166, 167. Left palp, prolateral (166) and retrolateral (167) views; arrow: femur apophysis. 168, 169. Tip of left procursus, retrolateral (168) and prolateral (169) views. Scale lines: 0.5 mm (164), 0.2 mm (165±169). 2001 HUBER: PHOLCIDS OF AUSTRALIA 53

Figs. 170±172. Wugigarra wanjuru, male. 170. Chelicerae, frontal view. 171. Left palp, retrolateral view (note that the genital bulb is rotated out of its natural position); arrow: femur apophysis. 172. Left genital bulb (view not clear). Scale lines: 0.2 mm. of procursus and bulb (compare ®gs. 177 and as in ®g. 176, procursus and bulb as in ®gs. 184). 176±178. Legs light brown, dark rings on MALE (holotype): Total length 3.0, cara- femora (subdistally) and tibiae (proximally, pace width 1.39. Leg 1: 22.8 (6.0 ϩ 0.5 ϩ subdistally); tips of femora and tibiae whit- 6.3 ϩ 8.4 ϩ 1.6), tibia 2: 4.5, tibia 3: 3.1, ish; with curved hairs on all tibiae and meta- tibia 4: 4.0; tibia 1 l/d: 53. Habitus and pro- tarsi; without spines and vertical hairs; retro- soma shape as in ®gs. 173 and 174; carapace lateral trichobothrium of tibia 1 at 26%; tar- brown, slightly darker medially, with radial sus 1 with ϳ22 quite distinct pseudoseg- stripes and darker mark posteriorly around ments. Opisthosoma gray, with some ocular area. Eye pattern as in ®g. 174; dis- blackish spots except ventrally; genital plate tance PME-PME 0.185; diameter PME slightly darker, plate in front of spinnerets 0.105; distance PME-ALE 0.055; diameter hardly visible. AME 0.075. Clypeus brown, unmodi®ed; VARIATION: Tibia 1 in other male: 5.1. sternum ochre-brown, centrally lighter. Che- FEMALE: In general very similar to male, licerae brown, with two small cones on each but carapace higher with the two lobes al- side, with stridulatory ridges (®g. 175). Palps most touching each other. Opisthosoma fron- 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 173±175. Wugigarra nauo, male. 173. Habitus. 174. Prosoma, frontal view (slightly dorsal). 175. Chelicerae, frontal view. Scale lines: 1 mm (173, 174), 0.3 mm (175). todorsally apparently unmodi®ed. Chelicerae tralia, Australia; Sept. 26, 1995 (R. P. Mc- also with stridulatory ®les. Palpal distal seg- Millan), pitfall traps, in WAM (99/1781). ments enlarged (®g. 179). Tibia 1 in 3 fe- ETYMOLOGY: Named for the Kalamaia males: 4.5, 4.7, 5.3. Epigynum as in ®g. 180, (Kalamai is a valid short form), an aboriginal dorsal view as in ®g. 181. tribe in southern Western Australia. The spe- DISTRIBUTION: Known only from type lo- cies name is a noun in apposition. cality in South Australia (map 8). DIAGNOSIS: Distinguished from the closely MATERIAL EXAMINED: AUSTRALIA: related W. nauo by the much smaller size, by South Australia: Kirton Point, Port Lincoln: the apophyses on the male chelicerae (com- Male holotype above, with 1( 3& (SAM pare ®gs. 175 and 185), and by details of N1999/856±9). procursus and bulb (compare ®gs. 177 and 178 with 183 and 184). Wugigarra kalamai, new species MALE (holotype): Total length 1.7, cara- Figures 182±185 pace width 0.94. Leg 1: 17.3 (4.8 ϩ 0.4 ϩ 4.9 ϩ 6.1 ϩ 1.1), tibia 2: 2.9, tibia 3: 2.1, TYPE: Male holotype from Helena-Aurora tibia 4: 2.8; tibia 1 l/d: 61. Habitus and pro- Ranges (30Њ23ЈS, 119Њ38ЈE), Western Aus- soma shape very similar to W. nauo (cf. ®gs. 2001 HUBER: PHOLCIDS OF AUSTRALIA 55

Figs. 176±181. Wugigarra nauo. 176. Left male palp, retrolateral view; arrow: sperm duct opening. 177. Tip of left procursus, retrolateral view. 178. Left genital bulb, prolateral view. 179. Left female palp, retrolateral view. 180, 181. Epigynum, ventral (180) and dorsal (181) views. Scale lines: 0.5 mm (179), 0.3 mm (176, 178, 180, 181), 0.1 mm (177).

173 and 174); carapace ochre, posteriorly pair of large brown apophyses and two pairs with single dark spot. Ocular area slightly of smaller distal, black apophyses (®g. 185). darker; distance PME-PME 0.145; diameter Palps as in ®g. 182, procursus and bulb as in PME 0.075; distance PME-ALE 0.025; di- ®gs. 182±184. Legs light brown, without ameter AME 0.080. Clypeus and sternum rings; with curved hairs on tibia and meta- ochre. Chelicerae light brown, with proximal tarsus 1 only; without spines and vertical 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 182±185. Wugigarra kalamai, male. 182. Left palp, retrolateral view. 183. Left genital bulb, prolateral view. 184. Left procursus, retrolateral view. 185. Chelicerae, frontal view. Scale lines: 0.2 mm (182, 183, 185), 0.1 mm (184). hairs; retrolateral trichobothrium of tibia 1 at TRICHOCYCLUS SIMON, 1908 27%; tarsus 1 distally with ϳ8 quite distinct pseudosegments, proximally pseudosegmen- Trichocyclus Simon, 1908: 407 (type species by monotypy T. nigropunctatus Simon, 1908; ex- tation not visible in dissecting microscope. amined).ÐDeeleman-Reinhold, 1993: 325; Opisthosoma shape as in W. nauo (cf. ®g. 1995: 35±36. 173); gray with many blackish spots except ventrally; genital plate brown. NOTE: The following diagnosis and de- FEMALE: Unknown. scription do not cover the ``aberrant'' T. wat- DISTRIBUTION: Known only from type lo- ta, which is assigned tentatively to Tricho- cality in southern Western Australia (map 8). cyclus. MATERIAL EXAMINED: AUSTRALIA: West- DIAGNOSIS: Small- to medium-sized (total ern Australia: Helena-Aurora Ranges: Male length usually 3±5 mm, rarely under 2 mm) holotype above. pholcids with globular or higher-than-long 2001 HUBER: PHOLCIDS OF AUSTRALIA 57 opisthosoma, mostly with large AME (AME (``a'' and ``p'' in ®gs. 197, 201, 206, 229, diameter usually 80±130% of PME diameter, 232, 260, 273; apophysis always distinct, in T. harveyi only 70%), apparently restricted pocket shallow to very deep), distally with to Australia or the Australian region. Distin- brush of hairlike membranous structures guished from Wugigarra (which is the only (e.g., ®gs. 216, 221, 229); bulb consisting of similar genus in Australia) by a characteristic proximal globular part and distal sclerotized weak zone dorsally on the male cymbium elements, often with transparent dorsal pro- (asterisks in ®gs. 201, 248, 265); by the ab- jection (arrows in ®gs. 191, 203, 252, 254, sence of the worm-shaped process on the 260; asterisk in ®g. 220); sperm duct opens bulb characteristic for Wugigarra; by the ab- close to this projection, without any embolus sence of curved hairs on the legs; by the (shafted arrow in ®g. 220). Tarsal organ cap- presence of several piriform gland spigots on sulate (®g. 208). Legs usually long (leg 1 the ALS (only 2 spigots on each spinneret in about 8±13 ϫ body length, in T. harveyi only Wugigarra); by the absence of stridulatory 5.6 ϫ body length), usually medium thin ®les in females; and by the characteristic and (tibia 1 l/d ϳ50±85, in T. harveyi only 34), conservative shape of the epigynum that has leg 1 always the longest, legs 2 and 4 about a median projection but lacks a pocket (e.g., same length, leg 3 shortest; with dark rings ®gs. 193, 213, 217). on femur subdistally, patella ϩ tibia proxi- DESCRIPTION: Total length in males usually mally, and tibia subdistally; tips of femora ϳ2.5±5 mm, only T. harveyi and T. ungumi and tibiae whitish; legs without spines and under 2 mm. Carapace oval, wider than long, curved hairs, with few vertical hairs; retro- with distinct thoracic groove not widened lateral trichobothrium of tibia 1 usually at 7± into thoracic pit, often with three pairs of lat- 16%, in T. harveyi at 30%; tarsus 1 some- eral spots loosely connected to median spot times with fairly distinct pseudosegments by radial marks (e.g., ®g. 187). Eight eyes in distally, sometimes pseudosegmentation not conservative pattern on moderately elevated visible in light microscope; in SEM regular ocular area. AME distinctively large (see pseudosegmentation visible (®g. 210). Opis- above), only in T. septentrionalis male on el- thosoma either globular or higher than long evation (®g. 244). Distance PME-ALE rela- (very ``high'' opisthosomata usually point in tively small (ϳ30±55% of PME diameter). dorsoposterior direction and could be regard- Clypeus never modi®ed. Male chelicerae ed as secondarily long; e.g., ®g. 204); gray usually with two pairs of distinctive frontal with black spots and sometimes white spots apophyses, the distal one more prominent dorsally; genital plate usually light brown, (e.g., ®gs. 190, 199, 211). Signi®cantly dif- about rectangular. Male gonopore without ferent cheliceral armature in T. septentrion- epiandrous spigots (®gs. 207, 219). ALS alis (®gs. 245, 246), arawari (®gs. 264, 266), with several piriform gland spigots (®gs. arnga (®gs. 269, 270), and bugai (®g. 274). 195, 209, 224, 225; examined: T. aranda, ar- Stridulatory ridges always present on male awari, arabana, nigropunctatus); other spin- chelicerae. Male palps large in relation to nerets typical for family. overall size (e.g., ®gs. 186, 275); coxa with- Sexual dimorphism slight, females with out retrolateral apophysis, trochanter without shorter legs, often with larger (higher) opis- apophyses; femur conspicuously enlarged thosoma; female chelicerae without stridula- (e.g., ®gs. 191, 238, 256), with retrolateral tory ridges. In most species female opistho- hump proximally, usually without ventral soma provided with two indistinct, usually apophyses (T. ungumi, T. harveyi, and T. whitish humps dorsofrontally; corresponding oborindi with brown ventral knob; ®g. 277); side on carapace either unmodi®ed or dis- patella triangular in lateral view (e.g., ®g. tinctively elevated (®g. 261). Epigynum 192); tibia simple, with 2 trichobothria in shape very conservative, consisting of two very proximal position (e.g., ®gs. 192, 206, plates, frontal plate with median elevation 247, 256); cymbium with characteristic weak (e.g., ®gs. 193, 213, 217), rarely with paired zone dorsally (e.g., ®gs. 201, 248, 265), with indentations (e.g., ®gs. 240, 241); internally heavily sclerotized procursus usually provid- simple, with pair of relatively small pore ed with dorsal apophysis and ventral pocket plates (e.g., ®gs. 194, 214, 234). 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

MONOPHYLY: All species included (except 13), and includes T. arawari, worora, arnga, T. watta) share the weak zone dorsally on the and bugai. These species share a distinctive male cymbium. This character is otherwise sclerotized element on the bulb set with only present in Aucana kaala, a ninetine scales (®gs. 259, 272), as well as relatively from New Caledonia. The transparent pro- huge pockets ventrally on the procursus; T. jection dorsally on the bulb might be a fur- arawari and worora have very similar pro- ther synapomorphy of the genus, but seems cursi, as do T. arnga and bugai. (7) T. harv- to be absent in some species (dif®cult to see). eyi and ungumi, both from Kimberley (map GENERIC RELATIONSHIPS: Trichocylus is ap- 14), have an unknown relation to each other parently part of holocnemines, whether this and to any other species. (8) Finally, T. watta group is mono- or paraphyletic. Within this is assigned tentatively to Trichocyclus. group, the presence of fairly distinct pseu- NATURAL HISTORY: The genus is most re- dosegments suggests it is basal, while the markable for its unparalleled ability among procursus with its dorsal apophysis and ven- pholcid spiders to thrive in the Australian de- tral pouch, as well as the reduction of epian- serts, in the ``dry heart'' of the continent. drous spigots, suggest it might form a clade Summarizing the information on collection with Physocyclus and Artema (and possibly with Wugigarra). Holocneminus might be labels, the spiders build their webs between close, but is insuf®ciently well known. See and under rocks, at rocky cliff faces and Relationships above (p. 6) for more detailed overhangs, at the bases of trees and under discussion. logs, and in caves, hollows and wells, and in SPECIFIC RELATIONSHIPS: Several species termite mounds and nests. No single-species groups can be identi®ed by morphological study, or any closer published observation, is characters and are roughly supported by their known to me. geographical distribution: (1) The nullarbor DISTRIBUTION: Presently known from Aus- group is most diverse in South Australia and tralia only. The CLD collection has a male southern Western Australia (map 9). T. nul- of an undescribed species from Togian Is- larbor and kokata have identical chelicerae; lands, Sulawesi, with remarkably similar T. nullarbor and hirsti have very similar pro- chelicerae but very different procursus. cursi. T. pandima is assigned to this group Within Australia, all described species are because of the long dorsal apophysis on the from the western part of the continent (west procursus, similar only to kokata. (2) The ni- of 140ЊE), with the remarkable exception of gropunctatus group, with three species in T. pustulatus. However, the collections stud- western Western Australia (T. nigropuncta- ied contain (apart from T. pustulatus) some tus, balladong, warianga) and with T. ara- unidenti®ed females from Queensland and bana widely distributed in central Australia New South Wales (map 2), suggesting that (map 10), share the shape of the procursus the genus is rare rather than absent in the and the wide but short distal apophyses on Great Artesian Basin and in the east. the male chelicerae. (3) The aranda group, COMPOSITION: The genus now includes a just to the north of the previous group (map total of 23 species. Several types of evidence 11), is distinguished by the rather slender suggest that the actual number is signi®cant- procursus (T. aranda, djauan, gnalooma). T. ly higher. First, nine species are known from kurara is assigned to this group because the male chelicerae are similar to those of T. just a single locality. Rather than indicating gnalooma). (4) T. septentrionalis is geo- limited distribution, this probably re¯ects the graphically close to the previous two groups, spotty effort made so far by collectors. Sec- but is highly autapomorphic in structure. (5) ond, the number of new species per decade The eastern group (T. grayi, oborindi, pus- represented in the collections studied does tulatus), which is presently the only group not seem to be leveling off; that is, six spe- known from Queensland (map 12), is distin- cies were represented until 1980, 14 species guished by the straight distal element of the until 1990, and 23 species until 2000. Third, procursus. (6) The arawari group, with four vast areas of Australia lack any record, prob- species, is only known from Kimberley (map ably in part because of collector bias. 2001 HUBER: PHOLCIDS OF AUSTRALIA 59

Trichocyclus nullarbor, new species dorsoposteriorly more elongated), chelicerae Figures 186±197 without stridulatory ridges. Tibia 1 in 11 fe- males: 6.8±10.3 (xÅ ϭ 8.2). Palpal tarsus tip TYPE: Male holotype from northern over- as in ®g. 196. Opisthosoma frontodorsally hang, Knowles Cave (31Њ08ЈS, 130Њ30ЈE), apparently without humps. Epigynum as in Nullarbor Plain, South Australia, Australia; ®g. 193, dorsal view as in ®g. 194. Several Sept. 29, 1988 (D. Hirst), in SAM (N1999/ piriform gland spigots on ALS (®g. 195). 842). DISTRIBUTION: Widely distributed in west- ETYMOLOGY: Named for the Nullarbor ern South Australia and southern Western Plain, where this species is common. The Australia (map 9). species name is a noun in apposition. MATERIAL EXAMINED: AUSTRALIA: South DIAGNOSIS: Easily distinguished from most Australia: Nullarbor Plain: Knowles Cave: congeners by the long narrow apophyses on Male holotype above, with 1& (SAM N1999/ the male chelicerae (®g. 190); from T. kokata 843); same data, 2( 5& several juveniles (which has identical chelicerae) by the shape (SAM N1999/844±50); Koonalda area of the dorsal apophysis on the procursus (31Њ24ЈS, 129Њ50ЈE), Feb. 1957 (C. Warner), (compare ®gs. 197 and 201); from T. hirsti 1( (SAM N1999/841); Koonalda Cave by the less voluminous procursus (compare (31Њ24ЈS, 129Њ50ЈE), Apr. 3, 1970 (J. Low- ®gs. 197 and 198), and by the apophyses on ry), 1( (WAM 93/2356); Warbla Cave the male chelicerae (compare ®gs. 190 and (31Њ31ЈS, 129Њ07ЈE), Oct. 10, 1966 (J. Low- 199). ry), 1& (WAM 93/2377); unnamed cave 8.6 MALE (holotype): Total length 3.7, cara- km E of Bore no. 5 off Murrawinjine Caves pace width 1.65. Leg 1: 31.8 (8.8 ϩ 0.7 ϩ Road, N of Nullarbor Station (ϳ31Њ10ЈS, 8.7 ϩ 11.7 ϩ 1.9), tibia 2: 6.3, tibia 3: 4.5, 131Њ00ЈE), Feb. 19, 1985 (N. Poulter), 1( 1 tibia 4: 6.2; tibia 1 l/d: 54. Habitus and pro- juvenile (AMS KS16911); unnamed cave soma shape as in ®gs. 186±189; carapace 14.5 km E of Bore no. 5 off Murrawinjine pale ochre with light brown pattern as in ®g. Caves Road, N of Nullarbor Station 187. Eye pattern as in ®g. 188; distance (ϳ31Њ10ЈS, 131Њ00ЈE), Feb. 18, 1985 (N. PME-PME 0.185; diameter PME 0.105; dis- Poulter), 1 juvenile assigned tentatively tance PME-ALE 0.045; diameter AME (AMS KS16909). Western Australia: Null- 0.135. Clypeus with wide, light brown band; arbor Plain: Top of Eucla Pass (31Њ39ЈS, sternum pale ochre, labium darker. Chelic- 128Њ52ЈE), Mar. 9, 1979 (collector not giv- erae ochre with pair of long, narrow frontal en), 1( 1 juvenile (AMS KS14997); un- apophyses, smaller cones at their bases, and named cave (31Њ11ЈS, 128Њ29ЈE), Sept. 14, stridulatory ridges (®g. 190). Palps as in ®gs. 1966 (J. Lowry), 1( 3& 3 juveniles (WAM 191 and 192, procursus as in ®g. 197. Legs 96/39±45); Old Homestead Cave (31Њ09ЈS, ochre, with indistinct darker rings on femora 127Њ56ЈE), Sept. 7, 1966 (J. Lowry), 1& (subdistally) and tibiae (proximally and sub- (WAM 93/2382), assigned tentatively; Skink distally), patellae also darker; tips of femora Hole (31Њ28ЈS, 127Њ55ЈE), Sept. 10, 1966 (J. and tibiae whitish; legs without spines, Lowry), 1( (WAM 93/2353); degraded do- curved, and vertical hairs; retrolateral tricho- line S of Mullamullang Cave (31Њ45ЈS, bothrium of tibia 1 at 15%; tarsal pseudo- 127Њ15ЈE), Jan. 8, 1966 (J. Lowry), 1& segments very indistinct, not countable. Op- (WAM 96/64); Kestral Cavern no.1 isthosoma ochre gray, dorsally with some (31Њ39ЈS, 127Њ13ЈE), cave doline, Jan. 9, blackish spots; genital plate light brown, 1972 (M. Gray), 1( (AMS KS56196); Fi- about square. restick Cave (31Њ46ЈS, 127Њ02ЈE), Oct. 17, VARIATION: Tibia 1 in 19 males: 6.5±10.4 1966 (J. Lowry), 1( (WAM 93/2379); Din- (xÅ ϭ 8.1). The specimens from near Perth go Cave (31Њ51ЈS, 126Њ44ЈE), Oct. 28, 1968 have slightly narrower dorsal apophyses on (J. Lowry), 3& 1 juvenile (WAM 93/2320± the procursus, and are therefore assigned ten- 3) assigned tentatively; Mullamullang Cave tatively. (31Њ44ЈS, 126Њ44ЈE), north doline, Jan. 3, FEMALE: In general very similar to male, 1966 (J. Lowry), 1( (WAM 93/ 2381); same but opisthosoma usually much higher (i.e., locality, doline, Aug. 30, 1966 (J. Lowry), 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 186±190. Trichocyclus nullarbor, male. 186. Habitus. 187±189. Prosoma, dorsal, frontal, and ventral views. 190. Chelicerae, frontal view. Scale lines: 1 mm (186±189), 0.3 mm (190). 2001 HUBER: PHOLCIDS OF AUSTRALIA 61

Figs. 191±194. Trichocyclus nullarbor. 191, 192. Left male palp, prolateral (191) and retrolateral (192) views; arrow: transparent bulbal process. 193, 194. Epigynum, ventral (193) and dorsal (194) views. Scale lines: 0.5 mm.

3& (WAM 92/2385±7); same locality, Jan. 6, ``et al.''), 1( 1& (WAM 99/1643±4); Cock- 1972 (M. Gray), 1& (AMS KS56198); Haig lebiddy Cave (31Њ58ЈS, 125Њ53ЈE), Jan. 10, Cave (30Њ44ЈS, 126Њ23ЈE), July 29, 1966 (J. 1966 (J. Lowry), 2( (WAM 93/2236±7); Lowry), 1( (WAM 93/2374); caves at Grass Patch (33Њ14ЈS, 121Њ44ЈE), house ceil- 30Њ51ЈS, 126Њ07ЈE, Apr. 15, 1990 (A. Baynes ing, Jan. 3, 1988 (A. F. Longbottom), 2( 1& 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 195, 196. Trichocyclus nullarbor, female. 195. ALS, showing several piriform gland spigots. 196. Tip of palp. Scale lines: 20 ␮m.

(WAM 99/1721±3); Great Victoria Desert, S 18, 1994 (D. Hirst/Pitlands Survey), in SAM of Skipper Knob (ϳ26Њ57ЈS, 126Њ20ЈE), wall (N1999/837). under overhang, Aug. 12, 1970 (J. Lowry), ETYMOLOGY: Named for the collector of 1( 2& (WAM 99/2112±4). The following this material and of many more pholcids in material from near Perth is assigned tenta- the South Australian Museum. tively (see above): Walyunga (31Њ44ЈS, DIAGNOSIS: Distinguished from the very 116Њ04ЈE), Nov. 28, 1981 (D. Hirst), 1( 1& similar T. nullarbor by the much stronger (SAM N1999/863±4); between Chittering dorsal apophysis on the procursus (compare and Pearce (ϳ31Њ33ЈS, 116Њ03ЈE), no date ®gs. 197 and 198) and by the apophyses on (G. H. Lowe), 1( (WAM 99/1638); Darling- the male chelicerae (distal apophyses wider, ton (31Њ54ЈS, 116Њ04ЈE), 1975±79 (G. H. proximal apophyses larger; compare ®gs. Lowe), 1( (WAM 99/1639); Gooseberry 190 and 199). Hill on Darling Swamp near Perth, Aug. 26, MALE (holotype): Total length 3.7, cara- 1971 (J. Lowry), 1( (AMS KS45158). pace width 1.84. Leg 1: 44.8 (11.9 ϩ 0.8 ϩ 11.7 ϩ 17.7 ϩ 2.7), tibia 2: 8.5, tibia 3: 6.0, Trichocyclus hirsti, new species tibia 4: 8.3; tibia 1 l/d: 68. Habitus and pro- Figures 198±200 soma shape as in T. nullarbor (cf. ®gs. 186± 189); carapace pale ochre with light brown TYPE: Male holotype from rocky cliff face pattern as in T. nullarbor, but less distinct. near ephemeral waterfall, 8 km SE of Mitch- Distance PME-PME 0.160; diameter PME ell Nob, Musgrave Ranges (26Њ11ЈS, 0.145; distance PME-ALE 0.045; diameter 131Њ53ЈE), South Australia, Australia; Oct. AME 0.120. Clypeus with wide, distally ta- 2001 HUBER: PHOLCIDS OF AUSTRALIA 63

Figs. 197±201. Trichocyclus nullarbor (197), T. hirsti (198±200), and T. kokata (201). 197, 198, 201. Left procursi, retrolaterodorsal views; ``a'', ``p'' ϭ apophysis and pocket; asterisk: weak zone on cymbium; unshafted arrow: more prominent in other male studied. 199. Male chelicerae, frontal view. 200. Epigynum, ventral view. Scale lines: 0.3 mm (197±199, 201), 0.5 mm (200). pering light brown band; sternum light imally, and stridulatory ridges (®g. 199). brown, with light speckles and larger spots Palps in general very similar to T. nullarbor near bases of coxae. Chelicerae ochre with (cf. ®gs. 191, 192), but procursus with much pair of strong, black apophyses distally and stronger dorsal apophysis (®g. 198). Legs another pair of shorter apophyses more prox- ochre-yellow, with indistinct darker rings on 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

roots was a necessary source over much of the area. The species name is a noun in ap- position. DIAGNOSIS: Easily distinguished from most congeners by the long narrow apophyses on the male chelicerae, which are identical to those of T. nullarbor (cf. ®g. 190); distin- guished from T. nullarbor by the long dorsal apophysis on the procursus (®g. 201). MALE (holotype): Total length 3.1, cara- pace width 1.35. Leg 1: 24.3 (6.7 ϩ 0.5 ϩ 6.7 ϩ 8.7 ϩ 1.7), tibia 2: 4.9, tibia 3: 3.3, tibia 4: 4.8; tibia 1 l/d: 50. Habitus and pro- soma shape as in T. nullarbor (cf. ®gs. 186± 189); carapace pale ochre with light brown Map 9. Distribution of Trichocyclus species: pattern similar to that of T. nullarbor (cf. ®g. nullarbor group. 187). Distance PME-PME 0.160; diameter PME 0.095; distance PME-ALE 0.045; di- ameter AME 0.120. Clypeus with wide, dis- femora (subdistally) and tibiae (proximally tally tapering light brown band; sternum pale and subdistally), patellae also darker; legs ochre, light brown speckled. Chelicerae iden- without spines, without curved and vertical tical to those of T. nullarbor (cf. ®g. 190), hairs; retrolateral trichobothrium of tibia 1 at ochre with pair of long, narrow frontal 10%; tarsus 1 distally with 13 quite distinct ϳ apophyses, smaller cones at their bases, and pseudosegments, proximally pseudosegmen- stridulatory ridges. Palps in general as in T. tation very indistinct. Opisthosoma ochre nullarbor (cf. ®gs. 191, 192), but procursus gray, covered with blackish spots except ven- with long dorsal apophysis (®g. 201). Legs trally; genital plate gray-brown, about rect- ochre, with darker rings on femora (subdis- angular. tally) and tibiae (proximally and subdistally), VARIATION: Tibia 1 in other male: 9.7; car- patellae also darker; tips of femora and tibiae apace width in this male: 1.61. whitish; legs without spines, without curved FEMALE: In general very similar to male; and vertical hairs; retrolateral trichobothrium tibia 1 in 2 females: 8.2, 8.8. Opisthosoma of tibia 1 at 12%; tarsal pseudosegments very frontodorsally apparently without humps. indistinct, hardly countable (apparently Epigynum as in ®g. 200, dorsal view not dis- Ͼ10). Opisthosoma roundish, ochre gray, tinguishable from T. nullarbor (cf. ®g. 194). dorsally with many blackish spots; genital ISTRIBUTION: Known only from type lo- D plate very light brown, about rectangular. cality in Musgrave Ranges, South Australia VARIATION: Tibia 1 in male from Mus- (map 9). grave Ranges: 8.1; this male is in general ATERIAL EXAMINED: AUSTRALIA: South M slightly larger, the pattern on the carapace is Australia: Musgrave Ranges: Male holotype more distinct, and the prolateral apophysis on above, with 1 2 (SAM N1999/838±40). ( & the procursus (unshafted arrow in ®g. 201) is slightly more prominent. Tibia 1 in males Trichocyclus kokata, new species from Kalamurina Station and Murda Hill: Figure 201 6.5, 7.1; these males differ slightly from the TYPE: Male holotype from Kolay Hut, Pa- holotype with respect to the procursus (dor- ney Station, Gawler Ranges (32Њ33ЈS, sodistal black ¯ap missing, dorsoproximal 135Њ36ЈE), South Australia, Australia; Dec. apophysis slightly more slender). 9, 1989 (D. Hirst), in SAM (N1999/832). FEMALE: In general very similar to male, ETYMOLOGY: Named for the Kokata, the but dark patterns on carapace and legs more so-called Gawler Range tribe, whose terri- distinct. Tibia 1 in 2 females accompanying tory included some of the most inhospitable male holotype: 7.2, 7.9. Opisthosoma fron- country in Australia; the water from tree todorsally apparently without humps. Epi- 2001 HUBER: PHOLCIDS OF AUSTRALIA 65 gynum externally not distinguishable from as in T. aranda (cf. ®g. 231), but femur with that of T. nullarbor (cf. ®g. 193). distinct dark knob ventrally subdistally; pro- DISTRIBUTION: Known from four localities cursus with long dorsal apophysis (®g. 203). in South Australia (map 9). Legs ochre-yellow, with barely visible darker MATERIAL EXAMINED: AUSTRALIA: South rings on femora (subdistally); tips of femora Australia: Gawler Ranges, Paney Station: and tibiae whitish; almost all hairs missing; Male holotype above, with 2& (SAM N1999/ retrolateral trichobothrium of tibia 1 at 10%; 833±4); Musgrave Ranges, Ngarutjara distally with ϳ12 quite distinct pseudoseg- (26Њ14ЈS, 131Њ47ЈE), amongst rocks, Oct. 13, ments, proximally pseudosegmentation very 1994 (D. Hirst), 1( 1 juvenile (SAM N1999/ indistinct. Opisthosoma roundish, gray, with 836); 1 km E of Rat Hole Yard, Kalamurina blackish and white spots except ventrally; Station (27Њ56ЈS, 138Њ00ЈE), Oct. 2±8, 1999 genital plate very light brown, about rectan- (D. Hirst), 1( (SAM NN9037); Simpson De- gular. sert, 1.7 km S of Murda Hill (26Њ58ЈS, VARIATION: Tibia 1 in other male: 11.7. 138Њ22ЈE), broad claypan between dunes, FEMALE: In general very similar to male, Mar. 24±29, 1998 (Sandy Desert Survey), but sternum darker, with many yellowish 1( (SAM NN9038). speckles and larger light spots near bases of coxae. Tibia 1 in 2 females: 12.8, 12.7. Op- Trichocyclus pandima, new species isthosoma frontodorsally with pair of dis- Figures 202, 203 tinct, sclerotized humps. Epigynum external- ly very similar to that of T. aranda (cf. ®g. TYPE: Male holotype from Dales Gorge, 233), but plate in front of frontal plate wider. Karijini National Park (22Њ28ЈS, 118Њ33ЈE), DISTRIBUTION: Known only from type lo- Western Australia, Australia; Sept. 11, 1981 cality, Karijini National Park, Western Aus- (D. Hirst), in SAM (N1999/798). tralia (map 9). ETYMOLOGY: Named for the Pandjima MATERIAL EXAMINED: AUSTRALIA: West- (also called Pand'ima), an aboriginal tribe in ern Australia: Karijini National Park, Dales Western Australia. The species name is a Gorge: Male holotype above, with 1( (and noun in apposition. a non-conspeci®c female) (SAM N1999/ DIAGNOSIS: Easily distinguished from most 799±800); same locality, July 24±25, 1998 congeners by the long dorsal apophysis on (D. Hirst), 1( 3& 5 juveniles (SAM N1999/ the procursus (®g. 203); from T. kokata 794±7). (which has a similar procursus; ®g. 201) by the shorter and wider distal cheliceral apoph- Trichocyclus arabana, new species yses and the shape of the bulb (®gs. 202, Figures 204±214 203). MALE (holotype): Total length ϳ3.5 (op- TYPE: Male holotype from 9.8 km WSW isthosoma shrunken), carapace width 2.03. of Mt. Bray, Mt. Barry Station (28Њ11ЈS, Leg 1: 14.9 ϩ 0.9 ϩ 15.1 ϩ 22.8; tarsus 134Њ42ЈE), South Australia, Australia; Sept. missing; tibia 2: 10.5, tibia 3: 6.8, tibia 4: 18±21, 1996 (Stony Desert Survey), in SAM 9.5; tibia 1 l/d: 81. Habitus and prosoma (N1999/814). shape as in T. nullarbor (cf. ®gs. 186±189); ETYMOLOGY: Named for the Arabana carapace pale ochre with light brown spot be- (Ngarabana), an aboriginal tribe of the Lake hind ocular area and three lateral spots hardly Eyre area, South Australia. The species name visible, without radial marks. Ocular area is a noun in apposition. with darker median and lateral marks; dis- DIAGNOSIS: Distinguished from the very tance PME-PME 0.160; diameter PME similar T. balladong by the much narrower 0.135; distance PME-ALE 0.030; diameter dorsal apophysis on the procursus (compare AME 0.135. Clypeus with slightly darker ®gs. 206 and 229); from all other congeners median band; sternum ochre-yellow, poste- by the short but wide distal apophyses on the riorly light brown with yellowish speckles. male chelicerae (®g. 211). Chelicerae with two pairs of apophyses and MALE (holotype): Total length 2.7, cara- stridulatory ridges (®g. 202). Palps in general pace width 1.35. Leg 1: 24.2 (6.8 ϩ 0.5 ϩ 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 202, 203. Trichocyclus pandima, male. 202. Chelicerae, frontal view. 203. Left procursus and genital bulb, retrolateral view; arrow: transparent bulbal process. Scale lines: 0.3 mm.

6.7 ϩ 8.9 ϩ 1.3), tibia 2: 4.7, tibia 3: 3.2, long (®g. 204), gray, covered with many tibia 4: 4.8; tibia 1 l/d: 50. Habitus as in ®g. black spots except ventrally. Genital plate 204. Prosoma shape as in T. nullarbor (cf. light brown; gonopore without epiandrous ®gs. 187±189); carapace pale ochre with spigots (®g. 207). Several piriform gland light brown pattern similar to that of T. nul- spigots on ALS (®g. 209). larbor (cf. ®g. 187), but without radial marks VARIATION: Tibia 1 in 24 males: 5.1±7.7 (xÅ and with only two pairs of lateral spots in- ϭ 6.6). In some specimens the pattern on the stead of three. Distance PME-PME 0.165; di- carapace resembles more that of T. nullar- ameter PME 0.095; distance PME-ALE bor, that is, there are radial marks and a third 0.035; diameter AME 0.105. Clypeus with pair of lateral spots; some specimens have a pair of brown bands converging distally; very long opisthosoma, but it is only the dor- sternum pale ochre, light brown speckled. sal part that is elongated, projecting far be- Chelicerae as in ®g. 211, ochre, with pair of yond the spinnerets. short, wide distal apophyses and proximal pair laterally, and stridulatory ridges. Palps FEMALE: In general very similar to male, as in ®gs. 205 and 206, procursus with but dark pattern on carapace more distinct, rounded dorsal apophysis (®g. 212). Legs and sternum darker (brown). Tibia 1 in 14 pale ochre, with light brown rings on femora females: 4.9±7.2 (xÅ ϭ 5.80). Palpal tarsal or- (subdistally) and tibiae (proximally and sub- gan as in ®g. 208. Pseudosegmentation of distally), patellae also darker; legs without tarsus 1 near tip as in ®g. 210. Opisthosoma spines, curved, and vertical hairs; retrolateral frontodorsally with pair of indistinct, trans- trichobothrium of tibia 1 at 12%; tarsal pseu- parent humps. Epigynum as in ®g. 213, dor- dosegments very indistinct, only distally ϳ7 sal view as in ®g. 214. discernible. Opisthosoma dorsoposteriorly DISTRIBUTION: Widely distributed in South 2001 HUBER: PHOLCIDS OF AUSTRALIA 67

Figs. 204±206. Trichocyclus arabana, male. 204. Habitus. 205, 206. Left palp, prolateral (205) and retrolateral (206) views; ``a'', ``p'' ϭ apophysis and pocket. Scale lines: 1 mm (204), 0.5 mm (205, 206).

Australia, southern Northern Territory, and (28Њ13ЈS, 134Њ32ЈE), Sept. 18±21, 1996 eastern Western Australia (map 10). (Stony Desert Survey), 1( (SAM N1999/ MATERIAL EXAMINED: AUSTRALIA: South 813); 5.5 km NW of Mt. Minyalcooroo, Australia: Mt. Barry Station: Holotype Arckaringa Station (27Њ57ЈS, 135Њ06ЈE), above; Jimmy Waterhole, Mt. Barry Station Sept. 15±20, 1996 (Stony Desert Survey), 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 207±210. Trichocyclus arabana. 207. Male gonopore. 208. Female palpal tarsal organ. 209. Male ALS, with several piriform gland spigots. 210. Female tarsus 1 near tip. Scale lines: 50 ␮m (207, 210), 10 ␮m (208, 209).

1( (SAM N1999/816); 31.5 km SE of Ucu- 131Њ21ЈE), Mar. 14±18, 1995 (Pitjantjatjara tanna Hill, Allendale Station (27Њ25ЈS, Lands Survey), 1( (SAM NN9022); 5.8 km 135Њ54ЈE), Nov. 14±17, 1995 (Stony Desert SSW of Mt. Goodair, Witjira National Park Survey), 1( (SAM N1999/817); sanddune (26Њ42ЈS, 135Њ36ЈE), Nov. 24, 1995 (Stony 4.5 km NE of Cheesman Peak (27Њ22ЈS, Desert Survey), 1( (SAM N1999/819); Ser- 130Њ42ЈE), Oct. 25, 1996 (Pitjantjatjara pentine Lake, west side on base of cliffs Lands Survey), 1( (SAM NN9021); 4.1 km (28Њ30ЈS, 129Њ00ЈE), Apr. 17, 1994 (D. ESE of Parke Camp Waterhole, Todmorden Hirst), 1( 5& (SAM N1999/823±8); quartz- Station (27Њ21ЈS, 134Њ29ЈE), Sept. 21±25, gibber plain 4.6 km ESE of Patsy Dam 1996 (Stony Desert Survey), 1( (SAM (28Њ37ЈS, 135Њ58ЈE), Oct. 26±31, 1995 (Lake N1999/818); 9.7 km S of Ampeinna Hills Eyre South Survey), 1( (SAM N1999/815); (27Њ09ЈS, 131Њ08ЈE), Mar. 19±23, 1995 (D. 4 km N of Halifax Hill (29Њ41ЈS, 135Њ49ЈE), Hirst; Pitlands Survey), 1( (SAM N1999/ Sept. 29±Oct. 5, 1995 (Painted Hills Sur- 820); 10.2 km E of Ampeinna Hills (27Њ05ЈS, vey), 1( (SAM N1999/812); Stuart High- 131Њ14ЈE), Mar. 19±23, 1995 (D. Hirst; Pit- way, roadside rest area (29Њ50ЈS, 135Њ08ЈE), lands Survey), 1( (SAM N1999/821); 14.2 May 11, 1999 (M. Gray, G. Milledge, H. km ESE of Maryinna Hill (27Њ00ЈS, Smith), 11&, 1 juvenile (AMS KS56195); 2001 HUBER: PHOLCIDS OF AUSTRALIA 69

Figs. 211±218. Trichocyclus arabana (211±214), T. nigropunctatus (215±218). 211, 215. Male che- licerae, frontal views. 212, 216. Left procursus, retrolaterodorsal views. 213, 217. Epigynum, ventral views. 214, 218. Epigynum, dorsal views. Scale lines: 0.5 mm (213, 214, 217, 218), 0.2 mm (211, 212, 215).

Stuart Highway, N of Port Augusta (31Њ15ЈS, Hirst), 1( (SAM N1999/810); Stone Dam, 136Њ32ЈE), May 10, 1999 (M. Gray, G. Mil- Paney Station, Gawler Ranges (32Њ35ЈS, ledge, H. Smith), 1( 1 juvenile (AMS 135Њ25ЈE), Dec. 11, 1989 (D. Hirst), 1( 2& KS56194); Hanker Caravan Park (31Њ53ЈS, (SAM N1999/802±4); Middleback Station 138Њ25ЈE), May 1, 1989 (D. C. Lee, D. (32Њ57ЈS, 137Њ23ЈE), Dec. 1983 (B. Guerin), 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

116Њ41ЈE), Western Australia, Australia; 1905 (W. Michaelsen), in MNHN (collection Simon 23021; examined). NOTES: As already noted by Deeleman- Reinhold (1993), the penultimate male in the Zoological Museum Hamburg that was col- lected at the same locality during the same expedition, and was identi®ed by Simon himself as T. nigropunctatus, is not a holo- type (contra Rack, 1961), but a paralecto- type. Unfortunately, I could not study any new material collected at the type locality. Be- cause females of Trichocyclus species are hardly or not at all distinguishable, the ma- Map 10. Distribution of Trichocyclus species: terial described herein is only tentatively nigropunctatus group. identi®ed as T. nigropunctatus, based pri- marily on geographic proximity. Yalgoo is 2( (SAM N1999/808±9). Western Australia: only 120 km from the nearest locality of the Warburton Mission (26Њ08ЈS, 126Њ25ЈE), in- material listed below, while it is 270 km side house, Aug. 25, 1970 (J. Lowry), 1( from the nearest locality of the next species 1& 1 juvenile (WAM 99/1960±2); Warri (T. balladong). Collecting males at and Well (24Њ53ЈS, 125Њ06ЈE), July 19, 1982 (B. around Yalgoo should easily solve this prob- Muir), 1( 1& (WAM 99/1735±6); Canning lem. Stock Route, Well 25 (22Њ59ЈS, 123Њ24ЈE), DIAGNOSIS: Distinguished from congeners inside dry well, July 31, 1987 (A. E. de by the distal male cheliceral apophyses that Jong), 2( 2& 1 juvenile (WAM 99/1626± appear divided into two parts each (®g. 215), 30); 7±8 km WNW of Point Salvation and by the tip of the procursus (®g. 216). (28Њ12ЈS, 123Њ36ЈE), Nov. 8±10, 1990, and MALE (Bush Bay): Total length 3.1, cara- Feb. 25±Mar. 5, 1991 (E. R. Pianka), 2( pace width 1.35. Leg 1: 26.0 (7.3 ϩ 0.5 ϩ (WAM 99/1660±1); 39 km E of Laverton 7.2 ϩ 9.5 ϩ 1.5), tibia 2: 5.2, tibia 3: 3.5, (28Њ28ЈS, 122Њ50ЈE), Nov. 1±4, 1989 (E. R. tibia 4: 5.9; tibia 1 l/d: 54. Prosoma shape, Pianka), 1( (WAM 99/1658); Red Sands colors, and pattern as in T. nullarbor (cf. ®gs. (28Њ12ЈS, 123Њ35ЈE), Nov. 5±7, 1989 (E. R. 186±189); ocular area with dark median Pianka), 1( (WAM 99/1662); Halls Creek mark posteriorly. Distance PME-PME 0.145; (18Њ14ЈS, 127Њ40ЈE), July 1, 1981 (D. Hirst), diameter PME 0.095; distance PME-ALE 1( (with a non-conspeci®c female!) (SAM 0.045; diameter AME 0.120. Clypeus with N1999/867±8). Northern Territory: West large brown mark tapering distally; sternum MacDonnell Natl. Park, Serpentine Gorge ochre, with brown speckles medially and (23Њ46ЈS, 132Њ59'), under rock, May 17, posteriorly. Chelicerae as in ®gs. 215 and 1999 (M. Gray, G. Milledge, H. Smith), 1( 223; ochre, with distinctive distal apophyses (AMS KS56178); Stuart Highway at that appear divided into long lateral and 21Њ38ЈS, 133Њ45ЈE, May 18, 1999 (M. Gray, shorter medial part. Palps in general as in T. G. Milledge, H. Smith), 1( 2& 1 juvenile arabana (cf. ®gs. 205, 206), only procursus (AMS KS65696). different, with much wider dorsal apophysis (®g. 216); with brush of hairlike structures Trichocyclus nigropunctatus Simon, 1908 (®g. 221). Bulb very similar to T. nullarbor Figures 215±225 (cf. ®gs. 191, 192); distal view as in ®g. 220. Legs ochre, with brown rings on femora Trichocyclus nigropunctatus Simon, 1908: 407.Ð (subdistally) and tibiae (proximally), patellae Deeleman-Reinhold, 1993: 325±327, ®gs. 1A±F. also darker; legs without spines, without TYPE: Female lectotype (designated by curved and vertical hairs; retrolateral tricho- Deeleman, 1993) from Yalgoo (28Њ21ЈS, bothrium of tibia 1 at 14%; tarsus 1 distally 2001 HUBER: PHOLCIDS OF AUSTRALIA 71 with ϳ9 barely distinguishable pseudoseg- at 26Њ12ЈS, 114Њ31ЈE, Aug. 22±Oct. 10, 1994 ments, proximally pseudosegmentation not (M. Harvey ``et al.''), 2( (WAM 99/2008± visible. Opisthosoma higher than long, but 9); same locality at 26Њ12ЈS, 114Њ25ЈE, Aug. posteriorly rounded, gray, covered with 22, 1994±Aug. 21, 1995 (N. McKenzie, J. many black and white spots except ventrally. Rolfe, P. West, N. Hall, M. S. Harvey), 9( Genital plate light brown; gonopore without 5& 3 juveniles (WAM 99/2010±25); same epiandrous spigots (®g. 219). Several piri- locality at 26Њ13ЈS, 114Њ35ЈE, Jan. 17±May form gland spigots on ALS (®g. 224). 17, 1995 (P. West ``et al.''), 3( 1& (WAM VARIATION: Tibia 1 in 80 males: 5.3±8.0 (xÅ 99/1989±92); same locality at 26Њ13ЈS, ϭ 6.7). 114Њ35ЈE, Jan. 12±Aug. 21, 1995 (N. Hall, P. FEMALE: In general very similar to male, West), 3( 1& 1 juvenile (WAM 99/1984±8); but sternum darker, and chelicerae without Francois Peron National Park (25Њ50ЈS, stridulatory ridges. Tibia 1 in 22 females: 113Њ36ЈE), Oct. 12, 1994±May 27, 1995 (M. 4.1±6.5 (xÅ ϭ 5.4). Palpal tarsus tip as in ®g. S. Harvey, N. McKenzie, J. Rolfe), 5( 1& 222. Opisthosoma frontodorsally apparently 1 juvenile (WAM 99/1900±6); Meedo Sta- without humps. Epigynum as in ®g. 217, dor- tion (25Њ43ЈS, 114Њ36ЈE), Aug. 22±Oct. 11, sal view as in ®g. 218. (Note that it is hardly 1994 (P. West ``et al.''), 2( 1 juvenile (WAM distinguishable from that of T. arabana; cf. 99/1949±51); same locality at 25Њ37ЈS, ®gs. 213, 214.) Spigots on ALS as in male 114Њ42ЈE, May 28±Aug. 21, 1995 (N. Hall), (®g. 225). 1( 1 juvenile, 1 prosoma (WAM 99/1940± DISTRIBUTION: Known from several local- 1); same locality at 25Њ41ЈS, 114Њ37ЈE, May ities in western Western Australia (map 10). 18±Aug. 22, 1995 (N. Hall), 1( (WAM 99/ MATERIAL EXAMINED: AUSTRALIA: West- 1948); same locality at 26Њ37ЈS, 114Њ41ЈE, ern Australia: Yalgoo: Female holotype May 16±21, 1995 (A. Sampey ``et al.''), 2& above; Greenough River, 2kmSWofEr- (WAM 99/1942±3), assigned tentatively; adu (28Њ43ЈS, 115Њ02ЈE), under rock, Feb. same locality at 25Њ39ЈS, 114Њ37ЈE, Aug. 22, 22, 1979 (M. R. Gray), 1( (AMS KS14914); 1994±Jan. 12, 1995 (P. West, N. McKenzie, Zuytdorp (27Њ16ЈS, 114Њ04ЈE), Aug. 26, J. Rolfe), 2& 3 juveniles (WAM 99/1944±7), 1994±May 18, 1995 (A. Sampey, M. S Har- assigned tentatively; Bidgemia Station, vey ``et al.''), 4( 5 juveniles (WAM 99/ Gascoyne Junction (25Њ07ЈS, 115Њ26ЈE), 2026±34); Nerren Nerren Station (27Њ00ЈS, Aug. 17, 1994±Aug. 20, 1995 (A. Sampey, 114Њ32ЈE), Aug. 25, 1994±Aug. 18, 1995 (P. N. Hall, J. M. Waldock), 3( 2 juveniles West, N. McKenzie, J. Rolfe, J. M. Waldock, (WAM 99/1848±52); same locality at N. Hall), 4( 2& 3 juveniles (WAM 99/ 25Њ10ЈS, 115Њ29ЈE, Jan. 16±June 4, 1995 (J. 1972±8); same locality at 27Њ03ЈS, 114Њ36ЈE, M. Waldock ``et al.''), 1& (WAM 99/1845) Aug. 25, 1994±May 28, 1995 (J. M. Wal- assigned tentatively; same locality at dock, P. West, N. McKenzie, J. Rolfe), 2( 25Њ05ЈS, 115Њ22ЈE, Aug. 17±Oct. 5, 1994 (A. 2& 1 juvenile (WAM 99/1979±83); Polelle Sampey ``et al.''), 1& (WAM 99/1853); same Station (26Њ55ЈS, 118Њ33ЈE), Aug. 5, 1982 locality at 25Њ11ЈS, 115Њ29ЈE, Aug. 17, (B. Y. Main), 1( (WAM 99/2102); Nanga 1994±Jan. 13, 1995 (N. McKenzie, J. Rolfe, Station (26Њ29ЈS, 114Њ03ЈE), Aug. 23, 1994± A. Sampey), 2( (WAM 99/1846±7); same Aug. 30, 1995 (A. Sampey, N. Hall, N. locality at 25Њ13ЈS, 115Њ31ЈE, Oct. 4, 1994± McKenzie, J. Rolfe, P. West), 11( 4& 4 ju- Jan. 13, 1995 (N. McKenzie, J. Rolfe), 1( 1 veniles (WAM 99/1952±71); Hamelin Pool juvenile (WAM 99/1841±2); same locality at (26Њ24ЈS, 114Њ10ЈE), Nov. 27, 1991 (M. S. 25Њ12ЈS, 115Њ30ЈE, Aug. 17±Oct. 4, 1994 (A. Harvey, M. E. Blosfelds), 1( 1 juvenile Sampey ``et al.''), 1& 1 juvenile (WAM 99/ (WAM 99/1645±6); Woodleigh Station 1843±4), assigned tentatively; same locality (26Њ12ЈS, 114Њ32ЈE), Oct. 11, 1994±May 17, at 25Њ03ЈS, 115Њ18ЈE, Jan. 13±June 6, 1995 1995 (P. West, N. McKenzie, J. Rolfe), 8( (J. M. Waldock ``et al.''), 2( 4 juveniles 1& 5 juveniles (WAM 99/1993±7, 1999± (WAM 99/1856±61); same locality at 2007); same locality at 26Њ11ЈS, 114Њ32ЈE, 25Њ03ЈS, 115Њ17ЈE, June 6±Aug. 20, 1995 May 17±Aug. 21, 1995 (N. Hall), 1& (WAM (N. Hall), 1& 1 juvenile (WAM 99/1862±3), 99/1998), assigned tentatively; same locality assigned tentatively; same locality, Aug. 17± 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 219±225. Trichocyclus nigropunctatus. 219. Male gonopore. 220. Genital bulb, distal view; shafted arrow: sperm duct opening; asterisk: transparent bulbal process. 221. Tip of procursus with brush of hairlike structures. 222. Tip of female palp. 223. Stridulatory ridges and apophyses on male right chelicera. 224, 225. ALS of male (224) and female (225), with several piriform gland spigots. Scale lines: 50 ␮m (219, 220, 223), 20 ␮m (221, 222, 224, 225). 2001 HUBER: PHOLCIDS OF AUSTRALIA 73

Oct. 5, 1994 (A. Sampey ``et al.''), 1& 2 ju- T. Ovtsharenko), 1( 1& (WAM 99/2092±5); veniles (WAM 99/1854±5), assigned tenta- same locality, WAPET camp (20Њ49ЈS, tively; Bush Bay (25Њ07ЈS, 113Њ48ЈE), Sept. 115Њ27ЈE), Nov. 5±Dec. 3, 1993 (M. S. Har- 27±Jan. 26, 1995 (P. West, M. S. Harvey, N. vey, J. M. Waldock), 1( 1 juvenile (WAM McKenzie, J. Rolfe), 3( 5& (WAM 99/ 99/1738); Barrow Island, Apr. 2, 1971 (A. A. 1877±84); Kennedy Range National Park Burbidge, W. H. Butler), 1( 2& (WAM 79/ (24Њ30ЈS, 115Њ01ЈE), Aug. 18, 1994±Aug. 334±6). 28, 1995 (N. McKenzie, J. Rolfe, M. S. Har- vey, P. West, N. Hall), 7( 5& 3 juveniles Trichocyclus warianga, new species (WAM 99/1911±24); same locality at Figures 226, 227 24Њ34ЈS, 114Њ57ЈE, Aug. 18±Oct. 8, 1994 (M. S. Harvey ``et al.''), 3( 1 juvenile TYPE: Male holotype from Barlee Range (WAM 99/1925±8); same locality at 24Њ30ЈS, Nature Reserve (23Њ23ЈS, 115Њ53ЈE), West- 115Њ02ЈE, May 29±Aug. 28, 1995 (N. Hall ern Australia, Australia; June 11±14, 1994 (P. ``et al.''), 2( (WAM 99/1908±9); Boola- G. & G. W. Kendrick) (WAM 99/1820). thana Station (24Њ25ЈS, 113Њ42ЈE), Sept. 27, ETYMOLOGY: Named for the Wariangga 1994±Aug. 25, 1995 (J. M. Waldock, N. (also Warianga), an aboriginal tribe in the Hall, N. McKenzie, J. Rolfe, A. Sampey), 6( Barlee Range area. The species name is a 1& 3 juveniles (WAM 99/1865±73, 1875); noun in apposition. same locality at 24Њ24ЈS, 113Њ40ЈE, May 28± DIAGNOSIS: Easily distinguished from sim- June 2, 1995 (J. M. Waldock ``et al.''), 1& ilar congeners (T. arabana, balladong, ni- (WAM 99/1864) assigned tentatively; same gropunctatus) by the presence of a dorsodis- locality at 24Њ24ЈS, 113Њ42ЈE, May 28±June tal apophysis on the procursus (arrow in ®g. 2, 1995 (J. M. Waldock ``et al.''), 1& (WAM 227). 99/1874) assigned tentatively; same locality MALE (holotype): Total length 2.9, cara- at 24Њ25ЈS, 113Њ46ЈE, Aug. 20±Sept. 30, pace width 1.19. Leg 1: 27.2 (7.6 ϩ 0.5 ϩ 1994 (A. Sampey ``et al.''), 1( (WAM 99/ 7.3 ϩ 10.1 ϩ 1.7), tibia 2: 5.1, tibia 3: 3.5, 1876); Mardathuna Station (24Њ24ЈS, tibia 4: 5.1; tibia 1 l/d: 62. Prosoma shape as 114Њ28ЈE), Oct. 5, 1994±May 28, 1995 (N. in T. nullarbor (cf. ®gs. 186±189); carapace McKenzie, J. Rolfe, A. Sampey), 5( (WAM ochre with wide brown median spot and 99/1933±7); same locality at 24Њ24ЈS, three pairs of lateral spots, without radial 114Њ27ЈE, Oct. 5, 1994±Jan. 14, 1995 (N. marks; ocular area posteriorly with median McKenzie, J. Rolfe), 1( 1& (WAM 99/ and lateral brown bands. Distance PME- 1938±9); same locality at 24Њ30ЈS, 114Њ38ЈE, PME 0.135; diameter PME 0.100; distance May 23±28, 1995 (A. Sampey ``et al.''), 1& PME-ALE 0.025; diameter AME 0.100. (WAM 99/1929), assigned tentatively; same Clypeus with pair of brown bands converg- locality at 24Њ27ЈS, 114Њ31ЈE, Oct. 5, 1994± ing distally; sternum light brown with yellow Jan. 14, 1995 (N. McKenzie, J. Rolfe), 1( 2 speckles, larger yellow spot behind labium. juveniles (WAM 99/1930±2); Cape Cuvier, Chelicerae as in ®g. 226; ochre, with short, Quobba Station (24Њ13ЈS, 113Њ30ЈE), Sept. wide apophyses distally, and smaller proxi- 29, 1994±May 30, 1995 (N. McKenzie, J. mal apophyses laterally; with stridulatory Rolfe, A. Sampey), 8( 5& 2 juveniles ridges. Palps in general as in T. arabana (cf. (WAM 99/1885±99); Barlee Range Nature ®gs. 205, 206), but procursus with distinctive Reserve (23Њ05ЈS, 115Њ47ЈE), June 15±18, dorsodistal apophysis (arrow in ®g. 227). 1994 (P. G. & G. W. Kendrick), 1( 3& Legs ochre-yellow, with light brown rings on (WAM 99/2139±41); same locality at femora (subdistally) and tibiae (proximally 23Њ03ЈS, 115Њ49ЈE, June 15±18, 1994 (P. G. and subdistally), patellae also darker, tips of & G. W. Kendrick), 2( 1& (WAM 99/2136± femora and tibiae whitish; legs without 8); Kuburu Well Cave, near Exmouth spines, curved, and vertical hairs; retrolateral (ϳ21Њ55ЈS, 114Њ07ЈE), Sept. 17, 1983 (J. trichobothrium of tibia 1 at 13%; tarsus 1 Lowry), 1( 2& (WAM 99/1716±18); Bar- distally with ϳ12 quite distinct pseudoseg- row Island, John Wayne Country (20Њ45ЈS, ments, proximally they are very indistinct. 115Њ22ЈE), Oct. 28, 1998 (M. S. Harvey, V. Opisthosoma dorsoposteriorly very long, 74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 226±229. Trichocyclus warianga (226, 227), and T. balladong (228, 229), male. 226, 228. Chelicerae, frontal views. 227, 229. Left procursi, retrolateral (slightly dorsal) views; unshafted arrow: diagnostic apophysis; ``a'', ``p'' ϭ apophysis and pocket. Scale lines: 0.3 mm. gray, covered with black and white spots ex- thosoma frontodorsally with pair of indis- cept ventrally; genital plate light brown, tinct, transparent humps. Epigynum external- about rectangular. ly not distinguishable from that of T. ara- VARIATION: Tibia 1 in 2 other males: 7.2, bana (cf. ®g. 213). 7.7. DISTRIBUTION: Known from two localities FEMALE: In general very similar to male, in western Western Australia (map 10). but sternum with median light band. Opis- MATERIAL EXAMINED: AUSTRALIA: West- 2001 HUBER: PHOLCIDS OF AUSTRALIA 75

ern Australia: Barlee Range Nature Reserve: VARIATION: Tibia 1 in 2 males from New- Male holotype above, with 1( 2& (WAM mans Rocks: 5.7, 5.7 (these males are in gen- 99/1821±3); Kennedy Range National Park eral much smaller that the holotype, but are (24Њ31ЈS, 114Њ58ЈE), Aug. 18±Oct. 4, 1994 identical in structure); tibia 1 in all other (M. S. Harvey ``et al.''), 1( (WAM 99/ males examined (N ϭ 5): 8.7±10.0 (xÅ ϭ 9.3). 1836). Some males have also white spots on the op- isthosoma. Trichocyclus balladong, new species FEMALE: In general very similar to male, Figures 228, 229 but sternum brown with yellowish speckles. Tibia 1 in 3 females from Newmans Rocks: TYPE: Male holotype from Tammin 4.3±4.9; in 2 females from Tammin: 7.7, 8.4. (31Њ38ЈS, 117Њ29ЈE), Western Australia, Aus- Opisthosoma frontodorsally with pair of in- tralia; May 1962 (B. Y. Main), in WAM (99/ distinct, transparent humps. Epigynum exter- 1698). nally not distinguishable from that of T. ar- ETYMOLOGY: Named for the Balardong abana (cf. ®g. 213). (also Balladong), an aboriginal tribe from the DISTRIBUTION: Known from several local- Tammin area. The species name is a noun in ities in southwestern Western Australia (map apposition. 10). DIAGNOSIS: Distinguished from the very MATERIAL EXAMINED: AUSTRALIA: West- similar T. arabana by the much wider dorsal ern Australia: Tammin: Holotype above, apophysis on the procursus (compare ®gs. with 1( 2& (WAM 99/1699±1701); same 206 and 229); from all other congeners by locality, Feb. 1963 (B. Y. Main), 1( 1& the short but wide distal apophyses on the (WAM 99/1686±7); same locality, 1963 (B. male chelicerae (®g. 228). Y. Main), 1( (WAM 99/1684); Perth airport MALE (holotype): Total length 3.7, cara- (31Њ58ЈS, 115Њ58ЈE), wet pitfalls, Jan. 6±Mar. pace width 1.77. Leg 1: 37.3 (10.1 ϩ 0.7 ϩ 18, 1994 (M. S. Harvey, J. M. Waldock), 2( 10.0 ϩ 14.4 ϩ 2.1), tibia 2: 7.3, tibia 3: 4.9, (WAM 99/2115±6); Mt. Elvire Station tibia 4: 7.5; tibia 1 l/d: 63. Prosoma shape as (29Њ19ЈS, 119Њ38ЈE), Sept. 13±17, 1994 (A. in T. nullarbor (cf. ®gs. 186±189); carapace Burbidge ``et al.''), 1( (WAM 99/1783); He- pale ochre with light brown pattern very sim- lena-Aurora Ranges (30Њ23ЈS, 119Њ38ЈE), ilar to that of T. nullarbor (cf. ®g. 187). Dis- Sept. 24, 1995 (R. P. McMillan), 1( (WAM tance PME-PME 0.215; diameter PME 99/1782); Kalgoorlie (30Њ48ЈS, 121Њ28ЈE), 0.120; distance PME-ALE 0.065; diameter Feb. 3±6, 1974 (T. Crawford), 1( (WAM 99/ AME 0.145. Clypeus with large brown mark 1570); Newmans Rocks (32Њ07ЈS, 123Њ10ЈE), tapering distally; sternum ochre-yellow with Feb. 24, 1989 (M. S. Harvey, M. E. Blos- triangular dark mark posteriorly. Chelicerae felds), 2(,3& (WAM 99/1728±32). as in ®g. 228; ochre, with pair of short, wide distal apophyses and smaller proximal pair, Trichocyclus aranda, new species with stridulatory ridges. Palps in general as Figures 230±234 in T. arabana (cf. ®gs. 205, 206), only pro- cursus different, with much wider dorsal TYPE: Male holotype from rocky cliff face, apophysis (®g. 229). Legs ochre, with light Glen Annie Gorge, Ruby Gap Nature Park brown rings on femora (subdistally) and (23Њ27ЈS, 135Њ00ЈE), Northern Territory, tibiae (proximally and subdistally), patellae Australia; Mar. 21, 1993 (D. Hirst), in SAM also darker, tips of femora and tibiae whitish; (N1999/786). legs without spines, curved, and vertical ETYMOLOGY: Named for the Aranda, an hairs; retrolateral trichobothrium of tibia 1 at aboriginal tribe from Northern Territory. The 13%; tarsus 1 distally with ϳ8 distinct pseu- species name is a noun in apposition. dosegments, proximally pseudosegmentation DIAGNOSIS: Distinguished from the very very indistinct. Opisthosoma dorsoposterior- similar T. djauan by the absence of two bul- ly longer than in T. nullarbor, ochre gray, bal apophyses (®g. 231; compare ®g. 235, covered with many black spots except ven- apophysis ``y'' and ®g. 236, arrow). From trally; genital plate light brown. other congeners by the apophyses on the 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 male chelicerae (small proximal and strong, Northern Territory and northwestern Western curved distal apophyses; ®g. 230). Australia (map 11). MALE (holotype): Total length 4.4, cara- MATERIAL EXAMINED: AUSTRALIA: pace width 1.87. Leg 1: 49.6 (13.6 ϩ 0.8 ϩ Northern Territory: Ruby Gap Nature Park: 13.5 ϩ 19.2 ϩ 2.5), tibia 2: 9.6, tibia 3: 6.4, Male holotype above, with 1( 3& 4 juve- tibia 4: 8.9; tibia 1 l/d: 78. Habitus and pro- niles (SAM N1999/787±90). All other ma- soma shape as in T. nullarbor (cf. ®gs. 186± terial is assigned tentatively. Western Austra- 189); carapace pale ochre with light brown lia: Woodstock Station, Site WS10 (21Њ40ЈS, pattern as in T. nullarbor, but radial spots 119Њ03ЈE), in cave, Oct. 26, 1990 (M. S. Har- less complete. Distance PME-PME 0.145; di- vey), 1( 1 juvenile (WAM 99/1834±5); ameter PME 0.145; distance PME-ALE Dales Gorge, Karijini National Park (22Њ28ЈS, 0.045; diameter AME 0.120. Clypeus with 118Њ33ЈE), Sept. 11, 1981 (D. Hirst), 1( 1& narrow brown band; sternum brown, with (SAM N1999/865±6); Barrow Island, cave ochre speckles. Chelicerae dark ochre with B4 (20Њ45ЈS, 115Њ22ЈE), Sept. 3, 1991 (W. F. pair of strong, curved black apophyses dis- Humphreys, B. Vine), 1( (WAM 99/1624). tally and another pair of small apophyses more proximally, and stridulatory ridges (®g. Trichocyclus djauan, new species 230). Palps as in ®g. 231, procursus as in ®g. Figures 235, 236 232. Legs ochre-yellow, with indistinct dark- er rings on femora (subdistally) and tibiae TYPE: Male holotype from Katherine (proximally and subdistally), patellae also Gorge (14Њ19ЈS, 132Њ28ЈE), Northern Terri- darker; legs without spines, curved, and ver- tory, Australia; Dec. 1980 (R. R. Jackson), in QMB (S34667). tical hairs; retrolateral trichobothrium of tibia ETYMOLOGY: Named for the Djauan, an ab- 1 at 13%; tarsus 1 distally with ϳ10 quite original tribe in the Katherine area of North- distinct pseudosegments, proximally pseu- ern Territory. The species name is a noun in dosegmentation not visible in dissecting mi- apposition. croscope. Opisthosoma roundish, gray, with DIAGNOSIS: Distinguished from the very many blackish spots except ventrally; genital similar T. aranda by the presence of two bul- plate gray-brown, about rectangular. bal apophyses (``y'' in ®g. 235 and arrow in VARIATION: Some specimens from Western ®g. 236). From other congeners by the Australia (see below) are very similar to the apophyses on the male chelicerae, identical type, but differ slightly with respect to the to those of T. aranda (small proximal and procursus (the proximal hump is more prom- strong, curved distal apophyses; cf. ®g. 230). inent: arrow ``1'' in ®g. 232; the tip is nar- MALE (holotype): Total length 3.6, cara- rower in dorsal view: arrow ``2'' in ®g. 232; pace width 1.55. Leg 1: 47.3 (12.0 ϩ 0.7 ϩ the distal ridge is smaller, and missing in the 12.3 ϩ 19.6 ϩ 2.7), tibia 2: 8.8, tibia 3: 5.7, male from Barrow Island: arrow ``3'' in ®g. tibia 4: 8.0; tibia 1 l/d: 77. Habitus and pro- 232), and the palpal femur which has a small soma shape as in T. nullarbor (cf. ®gs. 186± dark ventral hump subdistally. Also, PME- 189); carapace pale ochre with light brown PME distance is larger in these males (0.160) pattern as in T. nullarbor, but without radial while PME diameter is smaller (0.120). Tibia spots. Distance PME-PME 0.175; diameter 1 in the three known males from Western PME 0.120; distance PME-ALE 0.055; di- Australia: 12.7, 13.6, 14.1. More collecting ameter AME 0.145. Clypeus with large, dis- is needed to decide on the taxonomic status tally tapering brown band; sternum brown, of these specimens. with light spots anteriorly and smaller ones FEMALE: In general very similar to male; near bases of coxae. Chelicerae identical to tibia 1 in 3 females: 9.9, 10.5, 10.7. Opis- those of T. aranda (cf. ®g. 230). Palps also thosoma frontodorsally apparently without very similar to T. aranda (cf. ®g. 231), pro- humps. Epigynum as in ®g. 233, dorsal view cursus not distinguishable, but bulb signi®- as in ®g. 234. ALS with several piriform cantly different: with distinct proximal gland spigots. apophysis (unshafted arrow in ®g. 236) and DISTRIBUTION: Known from southern distal apophyses or crests (``x'' and ``y'' in 2001 HUBER: PHOLCIDS OF AUSTRALIA 77

Figs. 230±236. Trichocyclus aranda (230±234), T. djauan (235, 236). 230. Male chelicerae, frontal view. 231. Left male palp, retrolateral view. 232. Left procursus, retrolaterodorsal view; ``a'', ``p'' ϭ apophysis and pocket; arrows ``1±3'': variable structures (see text). 233, 234. Epigynum, ventral (233) and dorsal (234) views. 235, 236. Left procursus and genital bulb, retrolateral view (235), and left genital bulb, dorsal view (236); ``x'' and ``y'' ϭ diagnostic distal bulbal structures; unshafted arrow: diagnostic proximal bulbal apophysis. Scale lines: 0.5 mm (231, 233, 234), 0.3 mm (230, 232, 235, 236). 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

DISTRIBUTION: Known only from northern Northern Territory (map 11). MATERIAL EXAMINED: AUSTRALIA: Northern Territory: Katherine Gorge: Male ho- lotype above, with 2( 1& (QMB S50259); Kakadu National Park, Mirrai (Mt. Cahill), under stones (12Њ53ЈS, 132Њ42ЈE), July 5±7, 1994 (M. S. Harvey, D. M. Hyder), 1( 1& 1 juvenile (WAM 99/1807±9); Jabiru Resi- dency (12Њ41ЈS, 132Њ53ЈE), May 28, 1992 (M. S. Harvey), 1( (WAM 99/1673); 77 km N of Pine Creek (13Њ07ЈS, 131Њ50ЈE), under rocks, July 4, 1987 (M. S. Harvey), 1( (WAM 99/1676). The following material is assigned tentatively: Gregory National Park, Victoria Highway (15Њ36ЈS, 131Њ09ЈE), un- der rock, May 25, 1999 (M. Gray, G. Mil- ledge, H. Smith), 1( (AMS KS56177); Sad- Maps 11, 12. Distribution of Trichocyclus dle Creek, Victoria Highway (15Њ56ЈS, species: aranda group (map 11, top), and T. sep- 129Њ34ЈE), under logs, May 26, 1999 (M. tentrionalis and pustulatus group (map 12, bot- Gray, G. Milledge, H. Smith), 1( (AMS tom). KS56176).

Trichocyclus gnalooma, new species ®gs. 235, 236). Legs ochre-yellow, with Figures 237±241 darker rings on femora (subdistally) and tib- TYPE: Male holotype from Woodstock iae (proximally and subdistally), patellae also Homestead (21Њ37ЈS, 118Њ57ЈE), Western darker, tips of femora and tibiae whitish; legs Australia, Australia; July 28, 1987 (B. Y. without spines, without curved and vertical Main), inside house, in WAM (99/1792). hairs; retrolateral trichobothrium of tibia 1 at ETYMOLOGY: Named for the Ngaluma (also 10%; tarsus 1 distally with ϳ12 quite distinct Gnalooma), an aboriginal tribe in Western pseudosegments, proximally pseudosegmen- Australia. The species name is a noun in ap- tation not visible in dissecting microscope. position. Opisthosoma roundish, ochre-gray, with DIAGNOSIS: Distinguished from the similar blackish spots except ventrally; genital plate T. aranda by the shape of the procursus light brown, about rectangular. (compare ®gs. 232 and 239) and by the shape VARIATION: All specimens from other lo- of the distal male cheliceral apophyses (com- calities are signi®cantly smaller: tibia 1 in pare ®gs. 230 and 237). males from type locality: 11.1, 12.3; all other MALE (holotype): Total length 3.5, cara- localities: 5.6±8.5 (N ϭ 5). In addition, the pace width 1.84. Leg 1: 35.3 (9.7 ϩ 0.7 ϩ proximal apophysis on the genital bulb in 9.6 ϩ 13.2 ϩ 2.1), tibia 2: 7.1, tibia 3: 4.9, these males is less developed, and is even tibia 4: 6.9; tibia 1 l/d: 58. Prosoma shape as missing in the males from Saddle Creek and in T. nullarbor (cf. ®gs. 186±189); carapace Gregory National Park. The material from ochre with wide brown median band and the latter two localities is therefore assigned only two pairs of lateral spots, without radial tentatively. marks; ocular area brown, posteriorly with FEMALE: In general very similar to male; median and lateral darker bands. Distance tibia 1 in 2 females: 5.1 (Kakadu Natl. Park), PME-PME 0.175; diameter PME 0.120; dis- 11.3 (Katherine Gorge). Opisthosoma fron- tance PME-ALE 0.045; diameter AME todorsally with pair of indistinct, transparent 0.120. Clypeus with pair of brown bands humps. Epigynum externally not distinguish- converging distally; sternum brown with able from that of T. nullarbor (cf. ®g. 193). many yellowish speckles, larger spot near 2001 HUBER: PHOLCIDS OF AUSTRALIA 79

Figs. 237±241. Trichocyclus gnalooma. 237. Male chelicerae, frontal view. 238. Left male palp, retrolateral view. 239. Left cymbium and procursus, retrolaterodorsal view. 240, 241. Epigynum, ventral (240) and dorsal (241) views. Scale lines: 0.3 mm. coxae. Chelicerae as in ®g. 237; brown, with without curved and vertical hairs; retrolateral black apophyses, and stridulatory ridges. trichobothrium of tibia 1 at 12%; tarsus 1 Palps as in ®g. 238, procursus in retrolatero- distally with ϳ8 quite distinct pseudoseg- dorsal view with slender distal element (®g. ments, proximally they are very indistinct. 239). Legs ochre, with brown rings on fem- Opisthosoma dorsoposteriorly very long, ora (subdistally) and tibiae (proximally and gray, covered with black spots except ven- subdistally), patellae also darker, tips of fem- trally; genital plate brown, large, about ora and tibiae whitish; legs without spines, square. 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

VARIATION: Tibia 1 in 4 other males: 4.7 are similar to those of T. gnalooma (cf. ®g. (Boodarie Hill), 7.1, 7.1, 7.2. 237). FEMALE: In general very similar to male. MALE (holotype): Total length 3.4, cara- Opisthosoma frontodorsally with pair of in- pace width 1.74. Leg 1: 41.8 (10.9 ϩ 0.7 ϩ distinct, transparent humps. Epigynum as in 11.2 ϩ 16.5 ϩ 2.5), tibia 2: 7.9, tibia 3: 5.1, ®g. 240, with pair of distinctive indentations tibia 4: 7.2; tibia 1 l/d: 70. Habitus and pro- frontally; dorsal view as in ®g. 241. soma shape as in T. nullarbor (cf. ®gs. 186± DISTRIBUTION: Known from several local- 189); carapace ochre with three pairs of lat- ities in northwestern Western Australia (map eral spots; median and radial spots barely 11). visible; ocular area ochre. Distance PME- MATERIAL EXAMINED: AUSTRALIA: West- PME 0.145; diameter PME 0.125; distance ern Australia: Woodstock Homestead: Male PME-ALE 0.045; diameter AME 0.105. holotype above, with 2& (WAM 99/1793±4); Clypeus ochre; sternum pale ochre. Chelic- same locality, May 2, 1988 (J. M. Waldock), erae as in ®g. 242; brown, with black apoph- 1& (WAM 99/1670); Woodstock Station yses, and stridulatory ridges. Palps in general (21Њ37ЈS, 118Њ57ЈE), Feb. 10±17, 1989 (J. as in T. gnalooma (cf. ®g. 238), procursus Dell, R. A. How, J. M. Waldock), 1( (WAM and bulb as in ®g. 243. Legs ochre-yellow, 99/1832); same locality, Oct. 27, 1990 (M. with slightly darker rings on femora (subdis- S. Harvey), 1( (WAM 99/1831); same lo- tally) and tibiae (proximally and subdistally), cality at 21Њ37ЈS, 118Њ59ЈE, Feb. 10±17, patellae also darker, tips of femora and tibiae 1989 (J. Dell, R. A. How, J. M. Waldock), whitish; almost all hairs on legs missing; ret- 1( (WAM 99/1833); Boodarie Hill area, rolateral trichobothrium of tibia 1 at 11%; ϳ15 km SW of Port Hedland (ϳ20Њ25ЈS, tarsus 1 distally with ϳ7 fairly distinct pseu- 118Њ28ЈE), Oct. 11±17, 1994 (G. Harold, J. dosegments, proximally pseudosegmentation Dell), 1( (WAM 99/1741); Mundabullan- not visible in dissecting microscope. Opis- gana Station (20Њ32ЈS, 118Њ02ЈE), July 21, thosoma roundish, gray, with many dark 1981 (D. Hirst), 1( 2& (SAM N1999/869± spots dorsally; genital plate light brown, 71); Nigger Ring Rockhole, 7 km SSW of about rectangular. Camel Rock (22Њ37ЈS, 122Њ36ЈE), July 11, FEMALE: In general very similar to male, 1982 (B. Muir), 2& (WAM 99/1618±20), as- but opisthosoma much higher, though not signed tentatively; Lower Carawine Gorge pointed dorsoposteriorly. Tibia 1: 8.9. Opis- (21Њ29ЈS, 121Њ02ЈE), June 20±30, 1986 (K. thosoma frontodorsally with pair of fairly McKenna), 1& (WAM 99/1615), assigned distinct, transparent humps. Epigynum very tentatively. similar to that of T. gnalooma (cf. ®g. 240), but instead of frontal pockets only pair of sclerotized areas. Trichocyclus kurara, new species DISTRIBUTION: Known only from type lo- Figures 242, 243 cality in eastern Western Australia (map 11). MATERIAL EXAMINED: AUSTRALIA: West- TYPE: Male holotype from Glen Cum- mings Gorge (25Њ02ЈS, 128Њ18ЈE), Western ern Australia: Glen Cummings Gorge: Male Australia, Australia; Jan. 13±14, 1990 (M. S. holotype above, with 1& (WAM 99/1593). Harvey, T. F. Houston), in WAM (99/1592). ETYMOLOGY: Named for the Ngadadjara, Trichocyclus septentrionalis an aboriginal tribe from Western Australia, Deeleman-Reinhold, 1993 whose people of the Rawlinson Ranges are Figures 244±250 also called Kurara. The species name is a Trichocyclus septentrionalis Deeleman-Reinhold, noun in apposition. 1993: 327±328, ®gs. 1G±N. DIAGNOSIS: Distinguished from known congeners by the distinctively bifurcated bul- TYPE: Male holotype from outside cave bal tip (®g. 243) and by the shape of the C227 (22Њ03ЈS, 114Њ00ЈE), Cape Range, procursus (®g. 243); also by the shape of the Western Australia; May 24, 1990 (J. M. Wal- male cheliceral apophyses (®g. 242), which dock), in WAM (92/629), examined. (All 2001 HUBER: PHOLCIDS OF AUSTRALIA 81

Figs. 242, 243. Trichocyclus kurara, male. 242. Chelicerae, frontal view. 243. Left procursus and genital bulb, retrolateral view. Scale line: 0.3 mm. other specimens treated in the original de- in ®g. 248, with very broad but simple distal scription were designated paratypes.) part. Legs ochre, with indistinct darker rings DIAGNOSIS: Large species, easily distin- on femora and tibiae (subdistally), patellae guished from congeners by the huge protru- also darker; tips of femora and tibiae whitish; sions (and their shape) on the male chelicerae legs without spines, without curved and ver- (®gs. 245, 246), by the elevation in males tical hairs (most hairs missing in holotype); carrying the AME (®g. 244), by the procur- retrolateral trichobothrium of tibia 1 at 13%; sus with its wide distal part (®g. 248), and tarsal pseudosegments very indistinct, only by the large notch on the frontal plate of the distally ϳ10 visible. Opisthosoma rounded epigynum (®g. 249). as in T. nullarbor (cf. ®g. 186), ochre gray, MALE (holotype): Total length 4.2, cara- dorsally with some blackish spots; genital pace width 1.77. Leg 1: 50.1 (13.2 ϩ 0.9 ϩ plate light brown, about rectangular. 13.6 ϩ 19.6 ϩ 2.8), tibia 2: 10.0, tibia 3 VARIATION: Tibia 1 in 2 other males: 12.5, missing, tibia 4: 9.3; tibia 1 l/d: 73. Habitus 12.8. and prosoma shape as in T. nullarbor (cf. FEMALE: In general very similar to male, ®gs. 186±189), but AME on elevation (®g. but AME not on elevation, carapace laterally 244); carapace ochre, with brown spot be- with dark marks; sternum often darker than hind ocular area, without lateral spots. Eye in male, chelicerae without stridulatory ridg- pattern as in ®g. 244; distance PME-PME es. Tibia 1 in 16 females: 8.6±13.6 (xÅ ϭ 0.175; diameter PME 0.115; distance PME- 11.6); in one exceptionally small female: 5.6. ALE 0.055; diameter AME 0.120. Clypeus Opisthosoma frontodorsally without humps. light ochre-yellow; sternum ochre yellow, Epigynum as in ®g. 249, with distinctive with darker margins and slightly darker me- large notch; dorsal view as in ®g. 250. dian band. Chelicerae brown, with prominent DISTRIBUTION: Known only from Cape black apophyses and stridulatory ridges (®gs. Range, northwestern Western Australia (map 245, 246). Palps as in ®g. 247, procursus as 12). 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 244±250. Trichocyclus septentrionalis. 244. Male prosoma, dorsal view. 245, 246. Male che- licerae, frontal and lateral views. 247. Left male palp, retrolateral view. 248. Left cymbium and pro- cursus, retrolaterodorsal view; asterisk: weak zone on cymbium. 249, 250. Epigynum, ventral (249) and dorsal (250) views. Scale lines: 0.5 mm.

MATERIAL EXAMINED: AUSTRALIA: West- (WAM 99/1632); Cape Range, Shot Pot ern Australia: Cape Range, unnamed cave Cave, Sept. 21, 1983 (J. Lowry), 1& 1 ju- ``C94'', Sept. 16±17, 1983 (J. Lowry), 1& venile (WAM 99/1709±10); Cape Range, (WAM 99/1711); near Cape Range No. 2 Anomaly Cave ``C96'' near Learmonth Well, May 1965 (G. W. Kendrick, G. & (ϳ22Њ15ЈS, 114Њ05ЈE), Sept. 16±17, 1983 (J. T. Hitchin), 1& (WAM 99/1708); Cape Lowry), 2& (WAM 99/1712±5); Cape Range, July 8, 1987 (W. F. Humphreys), 1& Range, Camerons Cave ``C452'' (21Њ58ЈS, 2001 HUBER: PHOLCIDS OF AUSTRALIA 83

114Њ07ЈE), May 21, 1995 (J. M. Waldock), ®g. 186); ochre-gray, with many blackish entrance chamber, 1( 1& (WAM 99/1633± spots except ventrally; genital plate not dark- 4). er than surrounding area. Other than these specimens, I have seen FEMALE: Unknown. all the material listed in the original publi- DISTRIBUTION: Known only from type lo- cation (Deeleman-Reinhold, 1993), except cality in Northern Territory (map 12). for the three specimens deposited in the MATERIAL EXAMINED: AUSTRALIA: Rijksmuseum van Natuurlijke Historie, Lei- Northern Territory: Male holotype above. den, and two specimens in the WAM (92/ 131, 92/629). Trichocyclus oborindi, new species Figures 253, 254 Trichocyclus grayi, new species Figures 251, 252 TYPE: Male holotype from Bat Cave, Louie Creek (ϳ18Њ15ЈS, 138Њ05ЈE), Lawn TYPE: Male holotype from Stuart Highway Hill National Park, Queensland, Australia; at 21Њ37ЈS, 133Њ45ЈE, Northern Territory, Oct. 13, 1993 (R. Drysdale), in AMS Australia; May 18, 1999 (M. Gray, G. Mil- (KS37501). ledge, H. Smith), under rock, in AMS ETYMOLOGY: Named for the Ngoborindi (KS56192). (also called Oborindi), an aboriginal tribe ETYMOLOGY: Named for the ®rst collector from the Lawn Hill area, Queensland. The of the type specimen and of many more phol- species name is a noun in apposition. cids in the Australian Museum in Sydney. DIAGNOSIS: Distinguished from known DIAGNOSIS: Distinguished from known congeners by the shape of the procursus with congeners by the shape of the procursus with its small dorsal apophysis and long straight its distal prominent brush (®g. 252) and by distal element (®g. 254) and by the apophy- the male chelicerae with their rounded, wide ses on the male chelicerae (®g. 253; similar apophyses facing inwards (®g. 251). to T. grayi, ®g. 251). MALE (holotype): Total length 2.7, cara- MALE (holotype): Total length 3.9, cara- pace width 1.29. Leg 1: 25.1 (6.5 ϩ 0.5 ϩ pace width 1.77. Leg 1: 50.2 (13.3 ϩ 0.7 ϩ 6.8 ϩ 9.6 ϩ 1.7), tibia 2: 4.7, tibia 3: 3.2, 13.6 ϩ 20.3 ϩ 2.3), tibia 2: 8.8, tibia 3: 5.7, tibia 4: 4.6; tibia 1 l/d: 60. Habitus and pro- tibia 4: 8.0; tibia 1 l/d: 85. Habitus and pro- soma shape similar to T. nullarbor (cf. ®gs. soma shape similar to T. nullarbor (cf. ®gs. 186±189); carapace pale ochre with three 186±189); carapace ochre with small darker pairs of lateral spots with radial marks, with- spot behind ocular area and black line in tho- out median spot. Ocular area pale ochre; dis- racic furrow, without lateral and radial spots. tance PME-PME 0.105; diameter PME Ocular area light brown, slightly darker lat- 0.120; distance PME-ALE 0.040; diameter erally; distance PME-PME 0.135; diameter AME 0.105. Clypeus with small light spot, PME 0.135; distance PME-ALE 0.040; di- tapering distally; sternum whitish. Chelicerae ameter AME 0.105. Clypeus slightly darker ochre with two pairs of black, frontal apoph- than carapace; sternum brown with many yses, and stridulatory ridges (®g. 251). Palps yellowish speckles. Chelicerae ochre with in general as in T. harveyi (cf. ®g. 277), but two pairs of black, frontal apophyses, and femur ventrally without hump; procursus and stridulatory ridges (®g. 253). Palps in general bulb as in ®g. 252. Legs pale ochre, with as in T. harveyi (cf. ®g. 277), including dis- darker rings on femora (subdistally), patellae tinct ventrodistal hump on femur; procursus ϩ tibiae proximally, and tibiae subdistally; and bulb as in ®g. 254. Legs ochre-yellow, tips of femora and tibiae whitish; legs with- with slightly darker rings on femora (subdis- out spines, without curved and vertical hairs; tally), patellae ϩ tibiae proximally, and tibiae retrolateral trichobothrium of tibia 1 at 12%; subdistally; tips of femora and tibiae whitish; tarsus 1 distally with ϳ5 hardly visible pseu- legs without spines, without curved and ver- dosegments, proximally pseudosegmentation tical hairs; retrolateral trichobothrium of tibia not visible in dissecting microscope. Opis- 1 at 10%; tarsus 1 distally with ϳ13 quite thosoma shape similar to T. nullarbor (cf. distinct pseudosegments, proximally pseu- 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 251±254. Trichocyclus grayi (251, 252), and T. oborindi (253, 254), males. 251, 253. Chelic- erae, frontal views. 252, 254. Left procursi and genital bulbs, retrolateral views. Arrows: transparent bulbal processes. Scale lines: 0.3 mm. dosegmentation not visible in dissecting mi- mens from the Lawn Hill area, one of which croscope. Opisthosoma shape similar to T. is probably conspeci®c with the male holo- nullarbor (cf. ®g. 186); gray, with many type. The epigynum of this female is similar blackish spots dorsally; genital plate light to that of T. pustulatus (cf. ®g. 258), but is brown, about rectangular. medially more elevated; coloration as in FEMALE: The AMS has two female speci- male; tibia 1: 12.3. 2001 HUBER: PHOLCIDS OF AUSTRALIA 85

Figs. 255±258. Trichocyclus pustulatus. 255. Male chelicerae, frontal view. 256. Left male palp, retrolateral view. 257. Left cymbium and procursus, retrolaterodorsal view. 258. Epigynum, ventral view. Scale lines: 0.5 mm (256, 258), 0.3 mm (255, 257).

The other female has a dark pattern on the MATERIAL EXAMINED: AUSTRALIA: carapace similar to T. nullarbor (cf. ®g. 187), Queensland: Lawn Hill National Park: Male shorter legs (tibia 1: 7.6), and an even higher holotype above; Lawn Hill National Park, epigynum. Colonel Light Cave (ϳ18Њ15ЈS, 138Њ05ЈE), DISTRIBUTION: Known only from Lawn Hill Colless Ck., Oct. 28, 1993 (R. Drysdale), 1& National Park, western Queensland (map 12). (AMS KS37498), assigned tentatively. 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Trichocyclus pustulatus without humps. Tibia 1 in two females: 4.7, Deeleman-Reinhold, 1995 6.8. Figures 255±258 DISTRIBUTION: Known from several local- ities in northern Queensland (map 12). Trichocyclus pustulatus Deeleman-Reinhold, 1995: 36±37, ®gs. 9±15. MATERIAL EXAMINED: AUSTRALIA: Queensland: Mareeba: Male holotype and fe- TYPES: Male holotype and female paratype male paratype above; Chillagoe: female par- from Mareeba (17Њ00ЈS, 145Њ25ЈE), Queens- atype above; Chillagoe, Apr. 25, 1978 (R. land, Australia; July±Aug. 1992 (J. Wunder- Raven, V. E. Davies), 1& (QMB S50271), lich), in QMB (S51023±4), examined; and assigned tentatively; Wolfram (17Њ05ЈS, one additional female paratype from Chilla- 144Њ57ЈE), Feb. 13, 1972 (N. Clyde Cole- goe (17Њ09ЈS, 144Њ32ЈE), Queensland, Aug. man), 1& (QMB S50275), assigned tenta- 1992 (J. Wunderlich), under stone, in QMB tively; Almaden (17Њ21ЈS, 144Њ41ЈE), Mar. (S51025), examined. 1929 (W. D. Campbell), 1& (AMS KS65699), DIAGNOSIS: Similar to T. oborindi and T. assigned tentatively; Mt. Garnet (17Њ41ЈS, aranda, distinguished by the apophyses on 145Њ07ЈE), Feb. 24, 1972 (N. Clyde Cole- the male chelicerae (distal pair shorter, prox- man), 1& (QMB S50133), assigned tenta- imal pair more distal; ®g. 255), and by the tively. slightly different male cymbium and procur- sus (®g. 256). Trichocyclus arawari, new species MALE (holotype): Total length ϳ2.7 (op- Figures 259±266 isthosoma deformed), carapace width 1.48. Leg 1: 7.9 ϩ 0.6 ϩ 8.0, metatarsus and tarsus TYPE: Male holotype from Three-Mile missing, tibia 2: 5.7, tibia 3: 3.5, tibia 4 miss- Valley, Wyndham (ϳ15Њ30ЈS, 126Њ05ЈE), ing; tibia 1 l/d: 57. Habitus and prosoma Western Australia, Australia; June 30, 1981 shape similar to T. nullarbor (cf. ®gs. 186± (D. Hirst), in SAM (N1999/872). 189); carapace pale ochre with darker spot ETYMOLOGY: Named for the Ngarinjin behind ocular area and black line in thoracic (also called Arawari), an aboriginal tribe in furrow, with three pairs of light brown lateral Western Australia. The species name is a spots. Ocular area pale ochre with darker me- noun in apposition. dian line; distance PME-PME 0.175; diam- DIAGNOSIS: Distinguished from the very eter PME 0.120; distance PME-ALE 0.055; similar T. worora by the shape of the pro- diameter AME 0.135. Clypeus pale ochre cursus (compare ®gs. 265 and 268) and by with brown mark; sternum light brown with the shape of the male chelicerae that lack the many yellowish speckles. Chelicerae brown strong median projection (compare ®gs. 264 with black, frontal apophyses, and stridula- and 267); distinguished from all other known tory ridges (®g. 255). Palps as in ®g. 256, congeners also by the high elevations on the with small but distinct ventrodistal hump on female carapace (®g. 261). femur (smaller than in T. harveyi, cf. ®g. MALE (holotype): Total length 4.1, cara- 277); procursus as in ®g. 257. Legs light pace width 1.58. Leg 1: 36.9 (9.9 ϩ 0.7 ϩ ochre-yellow, with dark rings on femora 9.7 ϩ 14.1 ϩ 2.5), tibia 2: 6.9, tibia 3: 4.7, (subdistally), patellae ϩ tibiae proximally, tibia 4: 6.7; tibia 1 l/d: 63. Prosoma shape and tibiae subdistally; tips of femora and tib- and pattern as in T. nullarbor (cf. ®gs. 186± iae whitish; legs apparently without spines, 189), but without radial marks; ocular area without curved and vertical hairs (most hairs posteriorly with brown median band. Dis- missing); retrolateral trichobothrium of tibia tance PME-PME 0.145; diameter PME 1 at 15%. Opisthosoma deformed, but ap- 0.120; distance PME-ALE 0.045; diameter parently similar to T. nullarbor (cf. ®g. 186; AME 0.125. Clypeus with brown mark not see also original illustrations by Deeleman- reaching eyes; sternum brown with yellowish Reinhold, 1995), ochre-gray, with blackish speckles. Chelicerae as in ®g. 264; brown, and white spots dorsally. with only one pair of simple apophyses. FEMALE: Very similar to male; epigynum Palps as in ®gs. 259±260; procursus as in ®g. as in ®g. 258. Opisthosoma dorsofrontally 265. Legs ochre, with indistinct darker rings 2001 HUBER: PHOLCIDS OF AUSTRALIA 87

Figs. 259±263. Trichocyclus arawari. 259, 260. Left male palp, prolateral (259) and retrolateral (260) views; ``a'', ``p'' ϭ apophysis and pocket; arrow: transparent bulbal process. 261. Female prosoma, frontal view. 262, 263. Epigynum, ventral (262) and dorsal (263) views. Scale lines: 0.5 mm. on femora (subdistally) and tibiae (proximal- isthosoma dorsoposteriorly very long, gray, ly and subdistally); patellae also darker; most covered with black spots except ventrally; hairs on legs missing; retrolateral trichoboth- genital plate light brown, about rectangular. rium of tibia 1 at 14%; tarsus 1 distally with VARIATION: Tibia 1 in other males: 11.1, ϳ11 distinct pseudosegments, proximally the 11.7, 13.6, 13.7; in the males from Tunnel pseudosegmentation is very indistinct. Op- Creek National Park the prolaterodorsal 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 264±268. Trichocyclus arawari (264±266), and T. worora (267, 268), males. 264, 266, 267. Chelicerae, frontal views (264: male from Three-Mile Valley, Wyndham; 266: male from Napier Range). 265, 268. Left cymbia and procursi, retrolaterodorsal views; asterisk: weak zone on cymbium; arrows: prolaterodorsal apophysis. Scale lines: 0.3 mm (264±268). 2001 HUBER: PHOLCIDS OF AUSTRALIA 89

Trichocyclus worora, new species Figures 267, 268

TYPE: Male holotype from Kandiwal (Camp Creek) (14Њ52ЈS, 125Њ42ЈE), Mitchell Plateau, Western Australia, Australia; Dec. 11, 1993 (A. F. Longbottom), in WAM (99/ 1759). ETYMOLOGY: Named for the Worora, an aboriginal tribe in Kimberley, Western Aus- tralia. The species name is a noun in appo- sition. DIAGNOSIS: Distinguished from the very similar T. arawari by the shape of the pro- cursus (compare ®gs. 265 and 268) and by Maps 13, 14. Distribution of Trichocyclus the male chelicerae with their pair of strong species: arawari group (map 13, top); species of median projections (compare ®gs. 264 and unknown af®nities and T. watta (map 14, bottom). 267); from all other known congeners also by the high elevations on the female carapace (cf. ®g. 261). apophysis on the procursus (arrow in ®g. MALE (holotype): Total length 3.3, cara- 265) is simpler, more like that in T. worora pace width 1.52. Leg 1: 33.9 (9.2 ϩ 0.7 ϩ (cf. ®g. 268); the males from Geikie Range 9.0 ϩ 13.3 ϩ 1.7), tibia 2: 6.4, tibia 3: 4.4, and Napier Range have a posteriorly rounded tibia 4: 6.3; tibia 1 l/d: 64. Prosoma shape opisthosoma and lack both radial and lateral and pattern as in T. nullarbor (cf. ®gs. 186± marks on the carapace; the males from Na- 189), but without radial marks; ocular area pier Range differ signi®cantly with respect to and clypeus dark brown; sternum almost the chelicerae (®g. 266), but have apparently black. Distance PME-PME 0.145; diameter almost identical palpal structures. The spec- PME 0.120; distance PME-ALE 0.045; di- imens from Napier Range are therefore as- ameter AME 0.135. Chelicerae as in ®g. 267, signed tentatively. dark brown, with pair of simple distal apoph- FEMALE: In general very similar to male, yses, pair of strong median projections, and but carapace with pair of conspicuous ele- low lateral humps, all black. Palps in general vations (®g. 261). Opisthosoma frontodorsal- as in T. arawari (cf. ®gs. 259, 260), procur- ly with pair of indistinct, transparent humps sus as in ®g. 268. Legs light brown, with that oppose elevations on carapace. Tibia 1 black rings on femora (subdistally) and tibiae in 3 females: 10.1, 11.2, 11.3. Epigynum as (proximally and subdistally); patellae also in ®g. 262; dorsal view as in ®g. 263. darker, tips of femora and tibiae whitish; DISTRIBUTION: Known from several local- without spines, without curved and vertical ities in northern Western Australia (map 13). hairs; retrolateral trichobothrium of tibia 1 at MATERIAL EXAMINED: AUSTRALIA: West- 11%; pseudosegmentation of tarsi hardly vis- ern Australia: Three-Mile Valley, Wyndham: ible. Opisthosoma posteriorly more pointed Male holotype above, with 1( 2& (SAM than in T. nullarbor; gray, with black spots N1999/873±5); Napier Range, cave KN-90 except ventrally; genital plate dark brown. (17Њ14ЈS, 124Њ39ЈE), July 24, 1998 (S. M. VARIATION: Tibia 1 in male from Cockatoo Eberhard), 2( 1& 1 juvenile (WAM 99/ Island: 8.2. This male has a slightly more 2481±4), assigned tentatively; Tunnel Creek pointed prolaterodorsal apophysis on the pro- National Park (17Њ37ЈS, 125Њ10ЈE), June 7, cursus (arrow in ®g. 268), and the median 1999 (M. Gray, G. Milledge, H. Smith), in projection on the chelicerae is longer. webs, cave wall, 2( 1& (AMS KS56188); FEMALE: In general very similar to male, Geikie Range, cave KG-47 (18Њ02ЈS, but carapace with pair of conspicuous ele- 125Њ44ЈE), July 1, 1998 (S. M. Eberhard), vations (cf. ®g. 261), and opisthosoma dor- 1( (WAM 99/2473). soposteriorly much longer. Opisthosoma 90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 frontodorsally with pair of fairly distinct, visible in dissecting microscope. Opisthoso- transparent humps opposing elevations on ma roundish, but higher than long (2.7/2.4); carapace. Tibia 1 in 2 females: 6.5, 8.0. Epi- gray, with blackish spots except ventrally; gynum ventrally as in T. arawari (cf. ®g. genital plate not darker than surrounding 262). area. DISTRIBUTION: Known from two localities VARIATION: The other known male (see be- in northern Western Australia (map 13). low) is much smaller (carapace width: 1.00; Note, however, that the locality Cockatoo Is- tibia 1: 5.6), but the shapes of chelicerae and land is not certain. palpal structures appear identical. MATERIAL EXAMINED: AUSTRALIA: West- FEMALE: Unknown. ern Australia: Kandiwal (Camp Creek): DISTRIBUTION: Known from two localities Male holotype above, with 2& 1 juvenile in Kimberley, Western Australia (map 13). (WAM 99/1759±63); ?Cockatoo Island MATERIAL EXAMINED: AUSTRALIA: West- (16Њ06ЈS, 123Њ37ЈE), Oct. 30, 1961 (A. R. ern Australia: Jeremiah Hills: Male holotype Main), 1( (WAM 99/1693). above; Great Northern Highway, 52 km N of Turkey Creek (16Њ38ЈS, 128Њ12ЈE), June 7, Trichocyclus arnga, new species 1999 (M. Gray, G. Milledge, H. Smith), in Figures 269±272 webs, cave wall, 1( (AMS KS56182).

TYPE: Male holotype from Jeremiah Hills Trichocyclus bugai, new species (15Њ26ЈS, 128Њ44ЈE), Kimberley, Western Figures 273, 274 Australia, Australia; May 4, 1994 (R. D. Brooks), cave KJ-8, in WAM (99/2047). TYPE: Male holotype from Drysdale River ETYMOLOGY: Named for the Arnga, an ab- Station air®eld (15Њ43ЈS, 126Њ23ЈE), under original tribe in Kimberley, Western Austra- air®eld runway marker, Kimberley, Western lia. The species name is a noun in apposition. Australia, Australia; March 10, 1994 (A. F. DIAGNOSIS: Easily distinguished from Longbottom), in WAM (99/1750). known congeners by the huge median apoph- ETYMOLOGY: Named for the Bugai, aborig- yses on the male chelicerae (®gs. 269, 270). inal people of the Wenambal tribe in Kim- MALE (holotype): Total length 3.9, cara- berley, Western Australia. The species name pace width 1.65. Leg 1: 41.7 (11.3 ϩ 0.7 ϩ is a noun in apposition. 11.5 ϩ 16.1 ϩ 2.1), tibia 2: 8.3, tibia 3: 5.4, DIAGNOSIS: Distinguished from known tibia 4: 7.6; tibia 1 l/d: 78. Prosoma shape congeners by the single pair of cheliceral similar to T. nullarbor (cf. ®gs. 186±189); apophyses that are divided into two lobes carapace ochre with dark median band and each (®g. 274), and by the shape of the pro- wider spot behind ocular area, without lateral cursus (®g. 273; similar only to T. arnga, cf. marks. Ocular area dark brown; distance ®g. 271). PME-PME 0.175; diameter PME 0.120; dis- MALE (holotype): Total length 2.4, cara- tance PME-ALE 0.040; diameter AME pace width 1.03. Leg 1: 18.7 (5.2 ϩ 0.4 ϩ 0.120. Clypeus with large brown spot; ster- 5.1 ϩ 6.8 ϩ 1.2), tibia 2: 3.5, tibia 3: 2.4, num ochre with yellowish speckles. Chelic- tibia 4: 3.5; tibia 1 l/d: 55. Prosoma shape erae ochre with two very distinctive pairs of similar to T. nullarbor (cf. ®gs. 186±189); apophyses, as well as stridulatory ridges carapace ochre with pattern as in T. nullar- (®gs. 269, 270). Palps in general as in T. nul- bor (cf. ®g. 187). Ocular area with darker larbor (cf. ®gs. 191, 192), with distinctive median and lateral marks; distance PME- procursus (similar only to T. bugai) and bulb PME 0.120; diameter PME 0.080; distance (®gs. 271, 272). Legs ochre, with darker PME-ALE 0.040; diameter AME 0.105. rings on femora (subdistally) and patellae ϩ Clypeus with wide dark band tapering dis- tibiae proximally; legs without spines, with- tally; sternum ochre, medially with light out curved and vertical hairs; retrolateral tri- brown speckles. Chelicerae with only one chobothrium of tibia 1 at 7%; tarsus 1 dis- pair of bilobed apophyses, as well as strid- tally with ϳ20 quite distinct pseudoseg- ulatory ridges (®g. 274). Palps in general as ments, proximally pseudosegmentation not in T. arawari (cf. ®gs. 259, 260, rather than 2001 HUBER: PHOLCIDS OF AUSTRALIA 91

Figs. 269±274. Trichocyclus arnga (269±272), and T. bugai (273, 274), males. 269, 270, 274. Chelicerae, frontal and lateral views. 271, 273. Left procursi and genital bulbs, retrolateral views; ``a'', ``p'' ϭ apophysis and pocket. 272. Left genital bulb, prolateral view. Scale lines: 0.3 mm.

T. nullarbor!), with distinctive procursus 1 at 16%; tarsus 1 distally with ϳ15 very (similar only to T. arnga) and bulb (®g. 273). distinct pseudosegments, proximally pseu- Legs ochre, with darker rings on femora dosegmentation not visible in dissecting mi- (subdistally), patellae ϩ tibiae proximally, croscope. Opisthosoma probably roundish and tibiae distally; tips of femora and tibiae (damaged); gray, with blackish spots except whitish; legs without spines, curved, and ver- ventrally; genital plate dark brown, rectan- tical hairs; retrolateral trichobothrium of tibia gular. 92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

FEMALE: In general very similar to male, not visible in dissecting microscope. Opis- but sternum with wide brown median band; thosoma roundish (®g. 275), pale ochre-gray, tibia 1: 4.7. Opisthosoma frontodorsally with large blackish spots except ventrally; without humps. Epigynum in general similar genital plate not darker than surrounding to T. arabana (cf. ®g. 213) but with pair of area. distinctive, round invaginations near raised FEMALE: In general very similar to male; median part of frontal plate. tibia 1 in 2 females: 1.97, 2.03; carapace DISTRIBUTION: Known only from type lo- width in 2 females: 0.73, 0.77. Opisthosoma cality in Kimberley, Western Australia (map frontodorsally without humps. Epigynum as 13). in ®g. 281; dorsal view as in ®g. 282. MATERIAL EXAMINED: AUSTRALIA: West- DISTRIBUTION: Known only from type lo- ern Australia: Kimberley: Drysdale River cality in Kimberley, Western Australia (map Station air®eld: Male holotype above; same 14). collection data, 1& (WAM 99/1749). MATERIAL EXAMINED: AUSTRALIA: West- ern Australia: Kimberley, N of Larryoo: Trichocyclus harveyi, new species Male holotype above, with 2& 1 juvenile Figures 275±282 (WAM 99/1655±7).

TYPE: Male holotype from N of Larryoo Trichocyclus ungumi, new species (14Њ51ЈS, 126Њ49ЈE), Kimberley, Western Figures 283, 284 Australia, Australia; June 12, 1992 (M. S. Harvey, J. M. Waldock), under rock, in TYPE: Male holotype from 30 miles E of WAM (99/1654). Derby (17Њ19ЈS, 124Њ05ЈE), Kimberley, ETYMOLOGY: Named for the ®rst collector Western Australia, Australia; June 8, 1970 of the present material and of many more (Hemley Exped.), in WAM (99/1719). pholcids in the Western Australian Museum. ETYMOLOGY: Named for the Ongkomi DIAGNOSIS: Small species, distinguished (also called Ungumi), an aboriginal tribe from known congeners by the shape of the from Kimberley, Western Australia. The spe- procursus (®gs. 277, 280) and bulb (®gs. cies name is a noun in apposition. 277, 279), and by its small size and short legs DIAGNOSIS: Small species, distinguished (the legs of T. ungumi, the second smallest from congeners by the two-lobed distal male known species, are more than twice as long). cheliceral apophyses and the position of the MALE (holotype): Total length 1.4, cara- proximal apophyses (very proximal: ®g. pace width 0.74. Leg 1: 8.07 (2.16 ϩ 0.26 ϩ 283), and by the shape of the procursus 2.26 ϩ 2.68 ϩ 0.71), tibia 2: 1.52, tibia 3: (slightly similar to T. aranda, compare ®gs. 1.03, tibia 4: 1.65; tibia 1 l/d: 34. Habitus 232 and 284). and prosoma shape as in ®gs. 275 and 276; MALE (holotype): Total length 1.6, cara- carapace pale ochre with slightly darker pace width 0.97. Leg 1: 19.2 (5.2 ϩ 0.4 ϩ spots medially and laterally. Distance PME- 5.2 ϩ 7.2 ϩ 1.2), tibia 2: 3.5, tibia 3: 2.5, PME 0.075; diameter PME 0.080; distance tibia 4: 3.5; tibia 1 l/d: 52. Habitus and pro- PME-ALE 0.025; diameter AME 0.055. soma shape as in T. nullarbor (cf. ®gs. 186± Clypeus slightly darker than carapace; ster- 189), but much smaller; carapace ochre-yel- num pale ochre, shape as in T. nullarbor (cf. low with brown mark behind ocular area, ®g. 189). Chelicerae pale ochre with two black in thoracic groove, two pairs of lateral pairs of frontal apophyses, and stridulatory brown marks posteriorly, without radial ridges (®g. 278). Palps as in ®g. 277; femur marks. Ocular area with median dark band ventrally with distinct brown hump (®g. posteriorly. Distance PME-PME 0.095; di- 277); procursus as in ®g. 280, bulb as in ®g. ameter PME 0.085; distance PME-ALE 279. Legs ochre-gray, without rings; without 0.040; diameter AME 0.095. Clypeus with spines, without curved and vertical hairs; re- pair of brown bands under ALE, distally con- trolateral trichobothrium of tibia 1 at 30%; verging into large spot; sternum medially tarsus 1 distally with ϳ7 fairly visible pseu- light brown with yellowish spots, laterally dosegments, proximally pseudosegmentation ochre-yellow. Chelicerae ochre with pair of 2001 HUBER: PHOLCIDS OF AUSTRALIA 93

Figs. 275±278. Trichocyclus harveyi, male. 275. Habitus. 276. Prosoma, dorsal view. 277. Left palp, retrolateral view. 278. Chelicerae, frontal view. Scale lines: 0.5 mm (275±277), 0.1 mm (278). two-lobed distal apophyses and another pair ochre-yellow, with darker rings on femora very proximally, and stridulatory ridges (®g. (subdistally) and tibiae (proximally and sub- 283). Palps in general as in T. harveyi (cf. distally); patellae also darker; tips of femora ®g. 277), femur medioventrally with distinct and tibiae whitish; most hairs on legs miss- brown knob, bulb similar to that of T. aranda ing; retrolateral trichobothrium of tibia 1 at (cf. ®g. 231), procursus as in ®g. 284. Legs 14%; tarsus 1 distally with ϳ7 barely visible 94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 279±284. Trichocyclus harveyi (279±282), and T. ungumi (283, 284). 279. Left genital bulb, prolateral view. 280, 284. Left cymbia and procursi, retrolaterodorsal views. 281, 282. Epigynum, ventral (281) and dorsal (282) views. 283. Chelicerae, frontal view. Scale lines: 0.2 mm (279±284). pseudosegments, proximally pseudosegmen- FEMALE: In general very similar to male, tation not visible in dissecting microscope. with three pairs of lateral spots on carapace; Opisthosoma roundish, only slightly higher tibia 1: 4.5. Opisthosoma frontodorsally than long, ochre-gray, with many blackish without humps. Epigynum in ventral view as spots except ventrally; genital plate relatively in T. aranda (cf. ®g. 233). large, brown, trapezoidal. DISTRIBUTION: Known only from type lo- 2001 HUBER: PHOLCIDS OF AUSTRALIA 95 cality in Kimberley, Western Australia (map vial; same locality and collector, July 10, 14). 1979: 1& 1 juvenile (QMB S50176); Kakadu MATERIAL EXAMINED: AUSTRALIA: West- National Park, Mirrai (Mt. Cahill) (12Њ53ЈS, ern Australia: Kimberley, 30 miles E of Der- 132Њ42ЈE), July 5±7, 1994 (M. S. Harvey, D. by: Male holotype above, with 1& (WAM M. Hyder), under stones, 1& (WAM 99/ 99/1720). 1511), assigned tentatively.

Trichocyclus watta, new species MICROMERYS BRADLEY, 1877 Figures 285±292 Micromerys Bradley, 1877: 119 (type species by monotypy M. gracilis Bradley, 1877, exam- TYPE: Male holotype from Gorge NE of ined).ÐSimon, 1893: 474.ÐDeeleman-Rein- Mt. Gilruth (13Њ02ЈS, 133Њ05ЈE), Northern hold, 1986b: 205±224.ÐHuber, 1997c: 358. Territory, Australia; July 12, 1979 (G. B. Monteith), rainforest, sieved litter, in QMB DIAGNOSIS: Pale long-legged pholcids with (S50178). small ¯at prosoma and very long cylindrical ETYMOLOGY: Named for the Watta, an ab- opisthosoma (®g. 293); total length ϳ6±8 original tribe in the area of the Alligator Riv- mm; apparently restricted to the Australian ers, Northern Territory. The species name is region. Six eyes, AME either completely a noun in apposition. missing or reduced to pigment specks. Dis- DIAGNOSIS: Tiny species, easily distin- tinguished from Panjange (which is the only guished from all congeners and other possi- similar genus in Australia) by the single ble relatives (Wugigarra nauo, W. kalamai) sclerotized process on the bulb (e.g., ®gs. by the absence of AME. Also distinguished 302, 313, 316), the hinged process ventrally by the row of apophyses on the male chelic- on the procursus (``hp'' in ®gs. 296, 318), erae (®g. 288), and by the very short legs. the simple epigynum (e.g., ®gs. 303, 337; in MALE (holotype): Total length 1.2, cara- Panjange, the epigynum has a long scape), pace width 0.51. Leg 1: 3.50 (1.00 ϩ 0.19 ϩ the absence of eye turrets in the male, and 0.95 ϩ 0.97 ϩ 0.39), tibia 2: 0.84, tibia 3: the cylindrical opisthosoma. Distinguished 0.63, tibia 4: 0.95; tibia 1 l/d: 16. Habitus from Leptopholcus, Calapnita, and probably and prosoma shape as in ®gs. 285±287; also from Uthina (of which only females entire prosoma monochromous ochre. Ocular have been described) by the single bulbal area only slightly elevated (®g. 285); dis- projection and the hinged process on the pro- tance PME-PME 0.055; diameter PME cursus, and by the absence of frontal apoph- 0.055; distance PME-ALE 0.015; AME yses on the male chelicerae (®g. 299). missing. Sternum wide, similar to T. nullar- DESCRIPTION: Total length in males ϳ6±8 bor (cf. ®g. 189). Chelicerae light brown, mm. Carapace roundish to slightly longer with row of apophyses on each side than wide, ¯at and without markings. Six (®g. 288). Palps as in ®gs. 291 and 292, with eyes in two triads, sometimes slightly raised; weak zone dorsally on cymbium. Legs mon- AME missing or reduced to pigment specks ochromous ochre; without spines, without (e.g., ®g. 310). Distance PME-ALE small curved and vertical hairs; retrolateral tricho- (ϳ25±55% of PME diameter). Clypeus never bothrium of tibia 1 apparently at 58% (dif- modi®ed, in some species very short com- ®cult to see); tarsus 1 with ϳ5±7 pseudo- pared to all other pholcids (®gs. 294, 305, segments (dif®cult to count). Opisthosoma 310). Male chelicerae extremely conserva- oval, monochromous gray. tive, with pair of lateral apophyses, without FEMALE: Very similar to male. Chelicerae frontal apophyses (®g. 299), without stridu- apparently without stridulatory ®les. Epigyn- latory ridges. Male palps large in relation to um as in ®g. 289; dorsal view as in ®g. 290. prosoma size (®gs. 293); coxa without retro- DISTRIBUTION: Known from two localities lateral apophysis, trochanter with long ven- in Northern Territory (map 14). tral apophysis that is distally provided with MATERIAL EXAMINED: AUSTRALIA: distinct ridges (®gs. 300, 333); femur almost Northern Territory: NE of Mt. Gilruth: Male cylindrical, with small retrolateral hump holotype above, with 1& 4 juveniles in same proximally, with ventrodistal process in some 96 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 285±290. Trichocyclus watta. 285. Habitus, male. 286, 287. Male prosoma, frontal and dorsal views. 288. Male chelicerae, frontal view. 289, 290. Epigynum, ventral (289) and dorsal (290) views. Scale lines: 0.5 mm (285), 0.3 mm (286, 287, 289, 290), 0.1 mm (288). species (®g. 314); patella triangular in lateral 312, 318), and translucent fringed projection view; tibia considerably enlarged, with 2 tri- arising near base of hinged process (arrows chobothria (®g. 296); cymbium and procur- in ®gs. 301, 317, 320); bulb consisting ba- sus characteristically S-shaped (e.g., ®gs. sically of globular part and ¯at sclerotized 296, 312, 318; following dorsal contour); process (``p'' in ®g. 313); globular part of procursus consisting of main branch, com- bulb connected to cymbium by arched scler- plex hinged process ventrally (e.g., ®gs. 301, otized structure (``c'' in ®g. 313), to ¯at pro- 2001 HUBER: PHOLCIDS OF AUSTRALIA 97

Figs. 291, 292. Trichocyclus watta, left male pedipalp, prolateral (291) and retrolateral (292) views. Scale line: 0.2 mm. cess by triangular element (``t'' in ®g. 313). 315, 328), but mostly transparent and dif®- (I have not been able to ®nd the sperm duct cult to study. opening; the sperm duct may or may not en- NOTE: Deeleman-Reinhold (1986b) men- ter the ¯at sclerotized process.) Tarsal organ tioned a difference between the claws on tar- capsulate (®gs. 308, 330). Legs long and thin si 1±3 versus tarsus 4. A detailed reexami- (leg 1 ϳ4±5 ϫ body length, ϳ27±35 ϫ pro- nation of M. gracilis revealed no difference soma length; tibia 1 l/d ϳ75±85); leg formula between the claws on tarsi 1 and 4, except 1423, leg 4 signi®cantly longer than leg 2; that the claws on tarsus 4 were slightly lon- with dark ring in patella area and dark tibia- ger (32 versus 27 ␮m). metatarsus joint; legs without spines, without MONOPHYLY: All described species share vertical and curved hairs; retrolateral tricho- the bulbal morphology (¯at sclerotized pro- bothrium of tibia 1 at 2±4%; tarsi without cess, triangular element) and the single pair pseudosegmentation but with unique striation of lateral apophyses on the male chelicerae. of cuticle (®gs. 306, 332); tarsal claws as An undescribed species from Papua New usual for family (see Note below). Opistho- Guinea (see below) has the ``triangular'' el- soma very long and cylindrical, spinnerets at ement slightly more elaborate (longer, distal- the tip. Male gonopore with four epiandrous ly divided into two parts), but it is otherwise spigots (examined: M. daviesae, gracilis; ®g. very similar and clearly congeneric. The ®les 307). ALS with only one pair of spigots each on the tip of the male palpal trochanter (examined: M. daviesae, gracilis; ®g. 329); apophysis and the transparent projection aris- other spinnerets typical for family. ing from near the basis of the hinged process Sexual dimorphism slight, females with might be further synapomorphies of the ge- shorter legs, often with longer opisthosoma, nus. eye triads less raised. Epigynum shape ex- GENERIC RELATIONSHIPS: Two characters tremely simple and conservative; internally support the inclusion of Micromerys in the more diverse and complex (e.g., ®gs. 304, Pholcus group sensu Huber, 1995: the prox- 98 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 imolateral male cheliceral apophyses (func- leman-Reinhold (1986b), from Lake Boronto tionally related to the ventral trochanter near Somerset, 5 km S of Cape York apophysis); and the spindle-shaped male pal- (10Њ45ЈS, 142Њ35ЈE), Queensland, Australia; pal tibia. This was supported by cladistic Feb. 3±4, 1975 (R. Raven), in QMB (S885), analyses using equally weighted characters, examined. as well as by using successive weighting in REMARK: The selection of a female as type Hennig86. However, implied weighting in poses some problems as females of Microm- Pee-Wee suggested an alternative scenario at erys are extremely dif®cult to identify. How- all settings of the concavity constant except ever, M. gracilis seems to be the only species at conc ϭ 6: the South American Metagonia on Cape York Peninsula (map 15; note that was sister group of Micromerys, supported the locality Shiptons Flat in Deeleman-Rein- by the ventral hinged process on the procur- hold [1986b] rests on a misidenti®cation; see sus. (Both genera were nested in a ``Pholcus M. yidin below), suggesting that the material group sensu lato''.) studied is indeed conspeci®c with the neo- NATURAL HISTORY: The genus seems to be type. restricted to humid forests, where the spiders DIAGNOSIS: Distinguished from congeners live mainly on the underside of leaves (as by the wide main branch of the procursus judged from a few labels giving relevant in- (®g. 301) and by the straight bulbal apoph- formation; see also Deeleman-Reinhold and ysis that is slightly longer than the bulb (®gs. Deeleman, 1983, Deeleman-Reinhold, 1986a, 295, 302). b). No single-species study, or any closer ob- MALE (Murray Island): Total length 5.8, servation, is known to me. carapace width 0.84. Leg 1: 27.6 (7.3 ϩ 0.4 DISTRIBUTION: Known from Australia and ϩ 6.3 ϩ 10.9 ϩ 2.7), tibia 2: 5.1, tibia 3: (one undescribed species in the CLD collec- 3.2, tibia 4: 5.9; tibia 1 l/d: 79. Habitus and tion) from Papua New Guinea (Deeleman- prosoma shape as ®gs. 293, 294, 297, 298. Reinhold [1986a: ®g. 2; 1986b: 206] report- Entire prosoma whitish ochre, only proximal ed ``an unedited species from West Irian'', part of clypeus light brown (®g. 294). Ocular which I have not seen). Within Australia, the area ¯at; distance PME-PME 0.280; diameter genus is restricted to northern Northern Ter- PME 0.105; distance PME-ALE 0.050; AME ritory and to the Great Dividing Range in missing, only tiny black spot between triads. Queensland and New South Wales. It is re- Sternum slightly longer than wide (®g. 298). markable that four of the seven known spe- Chelicerae whitish, without modi®cation ex- cies occur within a limited area around cept pair of lateral apophyses (®g. 299). Cairns. Palps as in ®gs. 295 and 296, pale ochre, COMPOSITION: The genus now includes 7 only procursus and bulbal apophysis light to described species, 5 of them new. All de- dark brown; trochanter apophysis, procursus, scribed species are treated below. The col- and bulb as in ®gs. 300±302; tip of hinged lections studied contain numerous further vi- process as in ®g. 309; palpal tarsal organ as als with females only, which does not allow in ®g. 308. Legs pale ochre; patellae, femora useful guesses about possible undescribed tips, and tibia±metatarsus joints darker; with- species. For species previously assigned to out spines, without curved and vertical hairs; Micromerys, see Millot (1946), Brignoli retrolateral trichobothrium of tibia 1 at 2.3%; (1980), Deeleman-Reinhold (1986a), and tarsi without pseudosegmentation (®g. 306). Huber (1997c, 2000). Opisthosoma shape as in ®g. 293, monoch- romous pale ochre; two spigots on ALS (®g. Micromerys gracilis Bradley, 1877 307). Figures 293±309 VARIATION: Tibia 1 in 14 males: 6.1±7.4 (xÅ ϭ 6.7). Some males have no trace of AME, Micromerys gracilis Bradley, 1877: 119.ÐSimon, others have black spots in that area, but lens- 1893: 474.ÐDeeleman-Reinhold, 1986b: 206± es are always missing. Some males have 209, ®gs. 5±9, 60 (female only; male see M. yidin below). darker smudges on the opisthosoma. FEMALE: In general very similar to male, TYPE: Female neotype designated by Dee- but clypeus proximally not darker. Tibia 1 in 2001 HUBER: PHOLCIDS OF AUSTRALIA 99

Figs. 293±296. Micromerys gracilis, male. 293. Habitus. 294. Prosoma, frontal view. 295, 296. Left palp, prolateral (295) and retrolateral (296) views; ``hp'' ϭ hinged process. Scale lines: 1 mm (293), 0.5 mm (294±296).

9 females: 5.2±6.2 (xÅ ϭ 5.8). Tibia 1 in neo- DISTRIBUTION: Known from several local- type: 5.5. Variation in AME spots as in male ities in northern Northern Territory and (neotype without spots). Epigynum very sim- northern Queensland (map 15). ple, unsclerotized, with possibly distinctive MATERIAL EXAMINED: AUSTRALIA: arch shining through (®g. 303); dorsal view Queensland: Lake Boronto near Somerset: Fe- as in ®g. 304. male neotype above; Dividing Range, 15 km 100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 297±304. Micromerys gracilis. 297, 298. Male prosoma, dorsal and ventral views. 299. Male chelicerae, frontal view. 300. Tip of male palpal trochanter apophysis. 301. Left procursus, prolateral view; arrow: translucent fringed projection. 302. Left genital bulb, prolateral view. 303, 304. Epigynum, ventral (303) and dorsal (304) views. Scale lines: 0.4 mm (297, 298, 301±303), 0.2 mm (299, 304), 0.05 mm (300).

W of Captain Billy Creek (11Њ40ЈS, 141Њ50ЈE), Feb. 5±8, 1975 (G. B. Monteith), 142Њ45ЈE), July 4±9, 1975 (G. B. Monteith), rainforest, 1( (QMB S49738); Gordon 2 juveniles (QMB S49743), assigned tenta- Creek, Iron Range (12Њ43ЈS, 143Њ19ЈE), June tively; Andoom near Weipa (12Њ31ЈS, 24±30, 1976 (R. Raven, V. E. Davies), me- 2001 HUBER: PHOLCIDS OF AUSTRALIA 101

Figs. 305±309. Micromerys gracilis. 305. Male prosoma, frontal view. 306. Male tarsus 1 near tip, showing absence of pseudosegmentation. 307. Male ALS, with only two spigots. 308. Male palpal tarsal organ. 309. Tip of hinged process of right procursus, prolateral view. Scale lines: 300 ␮m (305), 100 ␮m (309), 30 ␮m (306, 308), 10 ␮m (307).

sophyll vine forest, 2& (QMB S886); same data, ``ϩ Lamond Hill'', 3& several juveniles (QMB S49741); West Claudie Range, Iron Range (12Њ45ЈS, 143Њ14ЈE), Dec. 3±10, 1985 (G. Monteith, D. Cook), rainforest, 50 m elev., 2( 3 juveniles (QMB S50306); same collection data, 2& 1 juvenile (QMB S50289); McIlwraith Range (ϳ13Њ43ЈS, 143Њ15ЈE), July 20, 1995 (J. Thompson, M. Moulds, T. Olive), 1( (AMS KS46098); Sil- ver Plains Station, near Chester R. (13Њ58ЈS, 143Њ22ЈE), Nov. 2, 1985 (I. Fanning) 1& (QMB S50307), assigned tentatively; Thurs- day Island, Torres Strait (10Њ35ЈS, 142Њ13ЈE), Aug. 1949 (N.L.H. Krauss), 1& in AMNH; Maps 15, 16. Distribution of Micromerys spe- Hammond Island, Torres Strait (10Њ32ЈS, cies in Australia. 142Њ13ЈE), July 4±8, 1974 (H. Heatwole, E. 102 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Cameron), 1& (QMB S50226), assigned ten- 5.9. Habitus and prosoma shape as in M. tatively; Murray Island, Torres Strait (9Њ56ЈS, gracilis (cf. ®gs. 293, 294, 297, 298). Entire 144Њ04ЈE), July 1974 (H. Heatwole, E. Cam- prosoma whitish ochre, only proximal part of eron), 2( 2& (QMB S50228); Yam Island, clypeus light brown. Distance PME-PME Torres Strait (9Њ53ЈS, 142Њ55ЈE), Nov. 28± 0.265; diameter PME 0.095; distance PME- Dec. 2, 1986 (J. Gallon), under bark, night ALE 0.035; AME missing, but distinct black coll., 1& (QMB S12400), assigned tentative- spots present (®g. 310). Chelicerae as in M. ly. Northern Territory: Gorge NE of Mt Gil- gracilis (cf. ®g. 299). Palps in general as in ruth (13Њ02ЈS, 133Њ05ЈE), July 10±13, 1979 M. gracilis (cf. ®gs. 295, 296), but femur (G. Monteith, D. Cook), 1& several juveniles ventrodistally with distinct apophysis (®g. (QMB S49771), assigned tentatively; Kaka- 314); procursus and bulb as in ®gs. 311±313. du National Park, Baroalba (12Њ50ЈS, Legs pale ochre, patellae (and femora tips) 132Њ57ЈE), July 6, 1994 (M. S. Harvey, D. and tibia±metatarsus joints darker; apparent- M. Hyder), 1& (WAM 99/1509), assigned ly without spines, without curved and verti- tentatively; Radon Creek (12Њ45ЈS, 132Њ53ЈE), cal hairs (many hairs missing); retrolateral rainforest, July 14±16, 1979 (G. Monteith, trichobothrium of tibia 1 at 2.0% (male in D. Cook), 2( 4& some juveniles (QMB SAM N1999/878); tarsi without pseudoseg- S49752); North Point, Kapalga (12Њ25ЈS, mentation. Opisthosoma shape as in M. grac- 132Њ22ЈE), July 19, 1979 (G. Monteith, D. ilis (cf. ®g. 293), monochromous pale ochre. Cook), 1( (QMB S49745); W Alligator, VARIATION: Tibia 1 in 8 males: 6.4±7.4 (xÅ ``mouth, WA2'' (12Њ11ЈS, 132Њ16ЈE), Nov. ϭ 6.8). 1979 (R. Raven), 1( 1& (QMB S49769); FEMALE: In general very similar to male, Kemp Airstrip (12Њ35ЈS, 131Њ20ЈE), rainfo- but clypeus proximally not darker. Tibia 1 (N rest, July 24±25, 1979 (G. Monteith, D. ϭ 3) 5.9±6.1. Epigynum very simple, unpig- Cook), 2( 1& 1 juvenile (QMB S49768); mented roundish plate; dorsal view as in ®g. same locality, Nov. 15±16, 1979 (R. Raven), 315. 2( 2& 2 juveniles (QMB S49755); Litch- DISTRIBUTION: Known only from the Cape ®eld National Park, Wangi Falls (13Њ09ЈS, Tribulation area and one suspicious locality 130Њ38ЈE), May 26, 1992 (M. S. Harvey, J. in New South Wales (map 15; see Note be- M. Waldock), 2( 1& 5 juveniles (WAM 99/ low). 1501±8). MATERIAL EXAMINED: AUSTRALIA: Queensland: Spear Creek: Male holotype Micromerys yidin, new species above, with 1( (QMB S49740); Cape Trib- Figures 310±315 ulation National Park, near beach (16Њ05ЈS, 145Њ28ЈE), July 24, 1992 (J. Coddington, G. Micromerys gracilis: Deeleman-Reinhold, 1986b: Hormiga), 3( 4& 2 juveniles (3 vials) 206±209, ®gs. 1±4 (male misidenti®ed). (USNM); Fritz Creek N of Bloom®eld TYPE: Male holotype from Spear Creek (15Њ52ЈS, 145Њ21ЈE), Dec. 1975 (M. Gray), (16Њ42ЈS, 145Њ24ЈE), Queensland, Australia; rainforest foliage, 1( 2& 5 juveniles (AMS Nov. 3±10, 1975 (R. Raven, V. E. Davies), KS457); Twelve Mile Scrub (15Њ50ЈS, in QMB (S34685). 145Њ19ЈE), Nov. 22±27, 1975 (collector not ETYMOLOGY: Named for the Idindji (also given), complex mesophyll vine forest, 1( 1 called Yidin), aboriginal rainforest dwellers juvenile (QMB S49739); Shiptons Flat from the Cairns area, northwestern Queens- (15Њ48ЈS, 145Њ15ЈE), Nov. 16±21, 1975 (R. land. The species name is a noun in apposi- Monroe, V. E. Davies), vine forest on basalt, tion. 2( 1& (QMB S887, 891); Home Rule, Mt DIAGNOSIS: Distinguished from congeners Hartley (15Њ44ЈS, 145Њ18ЈE), Nov. 11, 1974 by the shape of the lobes distally on the main (V. E. Davies, D.J.), on leaves, 2( 2& 1 ju- branch of the procursus (®gs. 311, 312), and venile (QMB S49742). New South Wales: by the very short, slightly curved bulbal Bruxner Park, Coffs Harbour (30Њ10ЈS, apophysis (®g. 313). 153Њ05ЈE), May 18, 1979 (D. Hirst), 1( MALE (holotype): Total length 6.4, cara- (SAM N1999/878), see Note below. pace width 0.71. Legs 1±3 missing, tibia 4: NOTE: The single record from New South 2001 HUBER: PHOLCIDS OF AUSTRALIA 103

Figs. 310±315. Micromerys yidin. 310. Male prosoma, frontal view. 311, 312. Left procursus, pro- lateral (311) and retrolateral (312) views. 313. Left genital bulb, prolateral view; ``c'' ϭ connecting piece between bulb and cymbium; ``t'' ϭ triangular element; ``p'' ϭ bulbal process. 314. Left male palpal trochanter and femur, retrolateral view. 315. Epigynum, dorsal view. Scale lines: 0.3 mm.

Wales might be based on some labeling error Australia; Nov. 22±27, 1975 (collector not and should be cited with some reservation given), complex mesophyll vine forest on until further material is collected from there. granite), in QMB (S49763). Revealingly, the single male shared the vial ETYMOLOGY: In Yidini, the aboriginal lan- with a Micromerys male of another species guage of the Cairns-Yarrabah region, gurran (M. gurran) that is also otherwise only means ``long''. This refers to the long bulbal known from the Cape Tribulation area. apophysis in this species. The species name is a noun in apposition. Micromerys gurran, new species DIAGNOSIS: Distinguished from congeners Figures 316±318 by the very long bulbal apophysis (®g. 316). TYPE: Male holotype from Twelve Mile M. wigi has a similar bulb, but the apophysis Scrub (15Њ50ЈS, 145Њ19ЈE), Queensland, is closer to the globular part, and the distal 104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 316±318. Micromerys gurran. 316. Left genital bulb, prolateral view. 317, 318. Left procursus, prolateral (317) and retrolateral (318) views; arrow: translucent fringed projection; ``hp'' ϭ hinged process. Scale line: 0.5 mm (316±318). lobes of the main branch of the procursus tibia±metatarsus joints darker; apparently differ in shape (compare ®gs. 316±318 with without spines, without curved and vertical 319±321). hairs (many hairs missing); retrolateral tri- MALE (holotype): Total length 8.1, cara- chobothrium of tibia 1 at 2.0%; tarsi without pace width 1.23. Leg 1: 33.2 (9.1 ϩ 0.5 ϩ pseudosegmentation. Opisthosoma shape as 8.0 ϩ 12.7 ϩ 2.9), tibia 2: 6.4, tibia 3: 4.1, in M. gracilis (cf. ®g. 293), monochromous tibia 4: 7.2. Habitus and prosoma shape as in ochre-gray. M. gracilis (cf. ®gs. 293, 294, 297, 298). VARIATION: Tibia 1 in 4 other males: 6.8, Carapace ochre to light brown, proximal part 7.2, 7.3, 7.5. of clypeus light brown; sternum whitish. Dis- FEMALE: Unknown. tance PME-PME 0.305; diameter PME DISTRIBUTION: Known only from the Cape 0.120; distance PME-ALE 0.045; AME with Tribulation area and one suspicious locality distinct black spots, but without lenses. Che- in New South Wales (map 15; see Note under licerae as in M. gracilis (cf. ®g. 299). Palps M. yidin description above). in general as in M. gracilis (cf. ®gs. 295, MATERIAL EXAMINED: AUSTRALIA: 296), but femur ventrodistally with distinct Queensland: Twelve Mile Scrub: Male holo- hump; procursus and bulb as in ®gs. 316± type above; Black Mtn (15Њ41ЈS, 145Њ13ЈE), 318. Legs pale ochre, coxae, patella area and Dec. 17, 1971 (N. Clyde Coleman), 2( 2001 HUBER: PHOLCIDS OF AUSTRALIA 105

(QMB S49746); Koah Road (16Њ49ЈS, DISTRIBUTION: Known only from type lo- 145Њ31ЈE), Apr. 2, 1972 (N. Clyde Coleman), cality, S of Cairns, Queensland (map 16). 1( (QMB S49750). New South Wales: Brux- MATERIAL EXAMINED: AUSTRALIA: ner Park, Coffs Harbour (30Њ10ЈS, 153Њ05ЈE), Queensland: Majors Mountain: Male holotype May 18, 1979 (D. Hirst), 1( (SAM N1999/ above, with 2 juveniles in same vial. 877), see Note under M. yidin description above. Micromerys gidil, new species Figures 322±324 Micromerys wigi, new species TYPE: Male holotype from Gordonvale Figures 319±321 (17Њ06ЈS, 145Њ47ЈE), Queensland, Australia; TYPE: Male holotype from Majors Moun- Feb. 1972 (N. Clyde Coleman), in QMB tain near Millaa Millaa (17Њ38ЈS, 145Њ32ЈE), (S49772). Queensland, Australia; Apr. 14±20, 1978 (R. ETYMOLOGY: In Yidini, the aboriginal lan- Raven, V. E. Davies), in QMB (S49753). guage of the Cairns-Yarrabah region, gidil means ``small''. This refers to the small size ETYMOLOGY: In Yidini, the aboriginal lan- guage of the Cairns-Yarrabah region, wigi of this species compared to the closest means ``big''. This refers to the larger size known relative. The species name is a noun in apposition. of this species compared to the closest DIAGNOSIS: Distinguished from congeners known relative. The species name is a noun by the shape of the bulbal apophysis with its in apposition. notch (arrow in ®g. 322) and by the shape of DIAGNOSIS: Distinguished from congeners the distal lobes of the main branch of the by the shape of the bulbal apophysis (®g. procursus (®gs. 323±324). M. wigi is very 319) and by the shape of the distal lobes of similar in shape but has relatively and ab- the main branch of the procursus (®gs. 320± solutely much larger palps (note that ®gs. 321). M. gidil is very similar in shape but 319±324 are all drawn to the same scale), has a notch on the bulbal apophysis, rela- and the PME are farther apart. tively and absolutely much smaller palps, MALE (holotype): Total length 6.1, cara- and the PME are closer together. pace width 0.87. Leg 1: 24.7 (6.5 ϩ 0.4 ϩ MALE (holotype): Total length 5.9, cara- 5.6 ϩ 9.7 ϩ 2.5), tibia 2: 4.4, tibia 3: 2.9, pace width ϳ1.0 (deformed). Leg 1: 28.0 tibia 4: 4.9. Habitus and prosoma shape sim- (7.3 ϩ 0.4 ϩ 6.7 ϩ 11.1 ϩ 2.5), tibia 2: 5.3, ilar to M. gracilis (cf. ®gs. 293, 294, 297, tibia 3: 3.5, tibia 4: 6.2. Habitus and prosoma 298), but carapace more roundish in dorsal shape similar to M. gracilis (cf. ®gs. 293, view. Carapace ochre, proximal part of clyp- 294, 297, 298), but carapace more roundish eus light brown; sternum whitish. Distance in dorsal view, as in M. gidil. Carapace PME-PME 0.185; diameter PME 0.105; dis- ochre, clypeus light brown except distal rim, tance PME-ALE 0.035; with distinct black sternum whitish. Distance PME-PME 0.305; spots in AME position, but without lenses. diameter PME 0.095; distance PME-ALE Chelicerae as in M. gracilis (cf. ®g. 299). 0.040; with distinct black spots at AME po- Palps in general as in M. gracilis (cf. ®gs. sition, but without lenses. Chelicerae as in M. 295, 296), but femur ventrodistally with dis- gracilis (cf. ®g. 299). Palps in general as in tinct bulge; procursus and bulb as in ®gs. M. gracilis (cf. ®gs. 295, 296), but femur 322±324. Legs pale ochre, patella area and ventrodistally with distinct bulge; procursus tibia±metatarsus joints darker; without and bulb as in ®gs. 319±321. Legs pale spines, without curved and vertical hairs; re- ochre, patella area and tibia±metatarsus joints trolateral trichobothrium of tibia 1 at 1.9%; darker; without spines, without curved and tarsi without pseudosegmentation. Opistho- vertical hairs; retrolateral trichobothrium of soma shape as in M. gracilis (cf. ®g. 293), tibia 1 at 4%; tarsi without pseudosegmen- pale ochre. tation. Opisthosoma shape as in M. gracilis VARIATION: Tibia 1 in 4 other males: 6.8, (cf. ®g. 293), pale ochre. 7.2, 7.3, 7.5. FEMALE: Unknown. FEMALE: Very similar to male. Tibia 1 in 106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 319±324. Micromerys wigi (319±321), and M. gidil (322±324). 319, 322. Left genital bulbs, prolateral views; unshafted arrow: diagnostic notch. 320, 323. Left procursi, prolateral views; shafted arrow: translucent fringed projection. 321, 324. Left procursi, retrolateral views. Scale line: 0.5 mm (319±324).

2 females: 5.9, 6.0. Epigynum extremely MATERIAL EXAMINED: AUSTRALIA: simple, unpigmented, roundish protruding Queensland: Gordonvale: Male holotype plate (similar to M. raveni, cf. ®g. 337); dor- above; Bellenden Ker Range, Cableway Base sal view similar to M. daviesae (cf. ®g. 328). Stn (17Њ16ЈS, 145Њ54ЈE), 100 m elev., Oct. DISTRIBUTION: Known only from the 17±24, 1981 (Earthwatch/QMB), 1( 1& 2 Cairns area, Queensland (map 16). juveniles (QMB S27754). 2001 HUBER: PHOLCIDS OF AUSTRALIA 107

Micromerys daviesae Deeleman-Reinhold, MATERIAL EXAMINED: AUSTRALIA: 1986 Queensland: Finch Hatton: Male holotype and Figures 325±333 female paratype above; Brandy Creek, E of Proserpine (20Њ21ЈS, 148Њ43ЈE), Apr. 24, Micromerys daviesae Deeleman-Reinhold, 1986b: 209±210, ®gs. 10±17, 24, 59d, 60. 1975 (R. Monroe, V. E. Davies), 1( (QMB S889); Rundle Range (23Њ29ЈS, 150Њ59ЈE), TYPE: Male holotype and female paratype on trees, Mar. 24±31, 1975 (R. Kohout, V. from Finch Hatton (21Њ09ЈS, 148Њ38ЈE), E. Davies), 3( 1& (QMB S49766); same Queensland, Australia; Apr. 10, 1975 (R. Ko- data, 1( 1& (QMB S890). hout, V. E. Davies), tangled web under leaf, in QMB (S888), examined. Micromerys raveni, new species DIAGNOSIS: Easily distinguished from con- Figures 334±338 geners by the single-pointed end of the main branch of the procursus (®gs. 326, 327). TYPE: Male holotype from Stott's Island MALE (Rundle Range): Total length 6.0, (28Њ14ЈS, 153Њ31ЈE), New South Wales, Aus- carapace width 0.97. Leg 1: 26.5 (6.8 ϩ 0.4 tralia; Nov. 18, 1978 (R. Raven), in QMB ϩ 6.0 ϩ 10.5 ϩ 2.8), tibia 2: 4.9, tibia 3: (S34684). 3.5, tibia 4: 5.6; tibia 1 l/d: 75. Habitus and ETYMOLOGY: Named for the collector of prosoma shape similar to M. gracilis (cf. ®gs. the type material and of many more pholcids 293, 294, 297, 298), but carapace more in the Queensland Museum in Brisbane. roundish in dorsal view. Entire prosoma pale DIAGNOSIS: Easily distinguished from con- ochre, proximal part of clypeus darker; ster- geners by the two pointed lobes distally on num whitish. Distance PME-PME 0.280; di- the main branch of the procursus (®gs. 335, ameter PME 0.105; distance PME-ALE 336). 0.035; no trace of AME. Chelicerae as in M. MALE (holotype): Total length 5.8, cara- gracilis (cf. ®g. 299). Palps in general as in pace width ϳ0.84 (deformed). Leg 1: 30.5 M. gracilis (cf. ®gs. 295, 296), but femur (7.9 ϩ 0.4 ϩ 7.2 ϩ 11.9 ϩ 3.1), tibia 2: 5.7, with ventrodistal bulge and procursus and tibia 3: 3.9, tibia 4: 6.5; tibia 1 l/d: 83. Hab- bulb distinctive (®gs. 325±327); tip of tro- itus and prosoma shape as in M. gracilis (cf. chanter apophysis as in ®g. 333; tip of ®gs. 293, 294, 297, 298). Entire prosoma hinged process as in ®g. 331; palpal tarsal pale ochre, proximal part of clypeus light organ as in ®g. 330. Legs pale ochre, patella brown; sternum whitish. Distance PME-PME area and tibia±metatarsus joints darker; ap- 0.280; diameter PME 0.100; distance PME- parently without spines, without curved and ALE 0.025; single black spot in area of vertical hairs; retrolateral trichobothrium of AME. Chelicerae as in M. gracilis (cf. ®g. tibia 1 at 3.5%; tarsi without pseudosegmen- 299). Palps in general as in M. gracilis (cf. tation (®g. 332). Opisthosoma shape as in M. ®gs. 295, 296), but femur ventrodistally with gracilis (cf. ®g. 293); monochromous pale rounded bulge; procursus, bulb, and trochan- ochre; gonopore with four epiandrous spigots ter apophysis as in ®gs. 334±336. Legs pale (®g. 329). ochre, patella area and tibia±metatarsus joints VARIATION: Tibia 1 in 3 other males: 5.7, darker; apparently without spines, curved, 6.7 (holotype), 6.9. (Note that the original and vertical hairs (many hairs missing); ret- measurements are mostly wrong, based on rolateral trichobothrium of tibia 1 at 2.0%; miscalibration and mismeasurement; e.g., tarsi without pseudosegmentation. Opistho- tibia 1 is 8.1 instead of 6.7, tibia 1 is shorter soma shape as in M. gracilis (cf. ®g. 293), than tibia 2; etc.) monochromous pale ochre. FEMALE: In general very similar to male, VARIATION: Tibia 1 in 2 other males: 6.8, but clypeus proximally not darker. Tibia 1 in 6.9. female paratype: 6.0. Epigynum very simple, FEMALE: In general very similar to male, unsclerotized, similar to M. raveni in ventral but clypeus proximally not darker. Tibia 1 in view (cf. ®g. 337); dorsal view as in ®g. 328. 2 females: 5.9, 6.0. Epigynum very simple, DISTRIBUTION: Widely distributed in mid- unpigmented roundish protruding plate (®g. dle-eastern Queensland (map 16). 337); dorsal view as in ®g. 338. 108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 325±328. Micromerys daviesae. 325. Left genital bulb, prolateral view. 326, 327. Left pro- cursus, prolateral (326) and retrolateral (327) views. 328. Epigynum, dorsal view. Scale lines: 0.3 mm (325±328).

DISTRIBUTION: Known from southeastern in pholcids. At ®rst, it might seem that this Queensland and northeastern New South simply results from Pholcus being the type Wales (map 16). genus, to which species were assigned before MATERIAL EXAMINED: AUSTRALIA: New the establishment of other genera, and to South Wales: Stott's Island: Male holotype which species tend to be assigned by workers above, with 1( 2& 2 juveniles (QMB unfamiliar with the other genera. Surprising- S49765). Queensland: Searys Scrub, Coo- ly, however, this seems not to be the case. loola (26Њ12ЈS, 153Њ03ЈE), Feb. 3, 1976 (R. Most of the ``old'' species not closely related Raven, V. E. Davies), 1( 1& 1 juvenile to the type species have been removed (58 (QMB S49767). species previously assigned to Pholcus have been synonymized or transferred to other PHOLCUS WALCKENAER, 1805 genera), and among the remaining 110 spe- REMARKS: With currently 110 nominal cies there are very few that are obviously species, Pholcus is by far the largest genus (e.g., P. everesti Hu and Li) or probably 2001 HUBER: PHOLCIDS OF AUSTRALIA 109

Figs. 329±333. Micromerys daviesae, male. 329. Gonopore, showing four epiandrous spigots. 330. Palpal tarsal organ. 331. Tip of hinged process of right procursus, prolateral view. 332. Male tarsus 1 near tip, showing absence of pseudosegmentation. 333. Tip of male palpal trochanter apophysis. Scale lines: 50 ␮m (329, 331), 20 ␮m (330, 332, 333).

(e.g., P. ciliatus Lawrence) misplaced. At the bolus lies between the uncus and appendix, same time, some species currently placed in is soft and transparent, and is thus easy to other genera might actually be Pholcus (e.g., overlook. Other characters, not exclusive for Spermophora elongata Yin and Wang; S. Pholcus, are the conservative male chelicerae longiventris Simon; S. faveauxi Lawrence). (a pair of dark frontal apophyses and a pair The characters most useful in distinguish- of light lateral apophyses; e.g., ®gs. 339, ing Pholcus from other genera seem to be the 369), the shape of the procursus (usually projections from the bulb, traditionally called with ventral boss; e.g., ®gs. 343, 366), and uncus, appendix, and embolus (``u'', ``a'', the knob- or worm-shaped apophysis on the and ``e'' in ®gs. 358, 372). The uncus is usu- often roughly triangular or oval epigynum. ally large, rather ¯at, heavily sclerotized, and Most Pholcus species further share a cy- is provided with many teeth or scales. The lindrical opisthosoma and eight eyes with lat- appendix is smaller, usually hook-shaped, eral triads and small AME. The latter char- also sclerotized, and is either a single rod or acter is especially interesting, as eye position split into two or even three parts. The em- has traditionally been a key character for dis- 110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 334±338. Micromerys raveni. 334. Left genital bulb, prolateral view. 335, 336. Left procursus, prolateral (335) and retrolateral (336) views. 337, 338. Epigynum, ventral (337) and dorsal (338) views. Scale lines: 0.3 mm (334±338). tinguishing pholcid genera. However, a re- simply because it lacks AME, but the geni- cent cladistic analysis (Huber, 2000) has sug- talia and the long opisthosoma suggest it is gested that AME have been lost many times also a six-eyed Pholcus. independently, and thus it comes as no sur- The real taxonomic problem in Pholcus is prise that this has also happened in Pholcus. its relationship to other genera of the Pholcus The new Australian species described below group, especially to Leptopholcus (Brignoli, as P. tagoman has the unmistakable genitalia 1980). This genus shares with Pholcus all de- of a real Pholcus, but it lacks any trace of tails of the chelicerae (including the modi®ed the AME. I strongly suspect that Spermopho- hairs imbedded in the frontal apophyses), has ra elongata was assigned to Spermophora a bulbal projection that seems to be a ho- 2001 HUBER: PHOLCIDS OF AUSTRALIA 111 mologue of the uncus, and has a soft trans- Therefore, Pholcus is currently represent- parent embolus, but apparently no appendix. ed in Australia by P. phalangioides and the Some Korean and northern Chinese Pholcus four species newly described below. Three of species seem indistinguishable in their geni- the endemic species occur in northern talia from Leptopholcus (e.g., most or all of Queensland and are closely related to each those described in Paik [1978] as well as P. other. The QMB has another closely related gaoi Song and Ren, and P. kwanaksaensis species, represented by a single poorly pre- Namkung and Kim). In fact, Leptopholcus served specimen from Horn Island, Torres might just be a polyphyletic group of pale, Strait. The six-eyed P. tagoman is restricted leaf-dwelling Pholcus species with a tenden- to the tropical north of Northern Territory, to cy to reduce the AME and the appendix. Kimberley in Western Australia, and possibly More relevant for the present work is the reaches into northwestern Queensland. Its presence of indigenous species of Pholcus in closest known relative is an undescribed spe- Australia. The only species previously re- cies (or maybe two species) from Malaysia corded (except P. phalangioides) was P. li- (in WAM 99/1994±5) and (interestingly) the toralis Koch, 1867. It is thus not surprising Seychelles (M. Saaristo, unpubl. MS). The that most Pholcus material in the collections male pholcid from Sumatra identi®ed by had been identi®ed as either P. phalangioides Kritscher (1957) (possibly correctly) as Sper- or P. litoralis. However, there is strong evi- mophora longiventris Simon also belongs in dence that P. litoralis is a synonym of P. this group. In fact, the Malaysian specimens phalangioides; that is, the illustrations in mentioned above might be ``Spermophora'' Koch (1872) clearly resemble P. phalangioi- longiventris. des rather than any of the species newly de- scribed herein, and at least two of the three Pholcus jinwum, new species putative syntypes in the ZMH are P. phal- Figures 339±353 angioides (one male from Sydney and one female without locality information; the third TYPE: Male holotype from of E Claudie specimen is a female from Queensland, but Scrub, Iron Range (12Њ43ЈS, 143Њ18ЈE), it lacks the opisthosoma; all were examined). Queensland, Australia; June 26, 1976 (col- All three vials carry labels presumably hand- lector unknown), in QMB (S34689). written by Koch himself, and the last two ETYMOLOGY: Named for the Jinwum, an could actually be types (the type localities aboriginal tribe in the Iron Range area, north- are Brisbane and Rockhampton). Therefore, ern Queensland. The species name is a noun P. litoralis Koch, 1867 is here synonymized in apposition. with P. phalangioides (Fuesslin, 1775) (NEW DIAGNOSIS: Similar to P. dungara, distin- SYNONYMY). guished by the shape of the procursus (com- A surprising fact is that P. ancoralis Koch, pare ®gs. 343 and 355) and by the dark pat- 1865, which is very widespread in the Paci®c tern on the epigynum (compare ®gs. 347 and region, is absent from the Australian collec- 356). tions studied. I have seen the type material of MALE (holotype): Total length 6.9, cara- this species (one male, two female prosomata pace width 1.9. Leg 1: 48.4 (12.0 ϩ 0.8 ϩ from Upolu, Samoa Islands, in the ZMH) and 11.9 ϩ 20.9 ϩ 2.8), tibia 2: 8.3, tibia 3: 6.1, numerous further specimens from the Samoa tibia 4: 7.7; tibia 1 l/d: 66. Prosoma shape as Islands (Sava'i), Fiji Islands, Moorea Islands in ®gs. 339 and 340. Carapace ochre, with (near Tahiti), New Hebrides (EspõÂritu Santo), brown mark broadly connecting to ocular and Loyalty Islands (UreÂa) (all in AMNH). In area (®g. 340); ocular area brown, clypeus contrast to the species newly described below, ochre, without marks; sternum ochre to light P. ancoralis is a representative of a mainly brown. Distance PME-PME 0.295; diameter southeast Asian species group characterized PME 0.160; distance PME-ALE 0.025; di- by a pair of horns between the eye triads (in- ameter AME 0.120. Chelicerae as in ®g. 339, cluding in addition at least P. bicornutus Si- with pair of black apophyses distally that are mon from the Philippines and P. dentifrons provided with two modi®ed hairs each (®g. Thorell from Burma). 341), and pair of unsclerotized rounded 112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 339±343. Pholcus jinwum, male. 339, 340. Prosoma, frontal and dorsal views. 341. Distal apophyses on chelicerae, with modi®ed hairs. 342, 343. Left palp, prolateral (342) and retrolateral (343) views. Scale lines. 1 mm (339, 340), 0.5 mm (342, 343), 0.05 mm (341). apophyses proximolaterally. Palps as in ®gs. tibia 1 at 4%; tarsal pseudosegments hardly 342 and 343; procursus as in ®g. 344. Tarsal visible in dissecting microscope, but regular organ capsulate. Legs ochre, distal ends of in SEM (®g. 351). Opisthosoma shape as in femora and tibiae whitish, without dark ®gs. 345 and 346; pale ochre, dorsally with rings; without spines, without curved and brown pattern as in ®g. 345, ventrally with vertical hairs; retrolateral trichobothrium of long brown line (®g. 346); gonopore in 2001 HUBER: PHOLCIDS OF AUSTRALIA 113

scanned male with ®ve epiandrous spigots MALE (holotype): Total length 6.8, cara- (®g. 350); several piriform gland spigots on pace width 1.8. Leg 1: 14.4 ϩ 0.9 ϩ 14.7, ALS (®g. 353). metatarsus and tarsus missing, tibia 2: 9.7, VARIATION: Tibia 1 in 8 males: 8.0±14.5 tibia 3: 6.7, tibia 4 missing; tibia 1 l/d: 79. (xÅ ϭ 11.4). In some specimens the proximal Habitus and prosoma shape as in P. jinwum cheliceral apophyses are slightly closer to- (cf. ®gs. 339, 340). Brown mark on carapace gether. similar to P. jinwum (cf. ®g. 340), but wider; FEMALE: In general very similar to male. ocular area brown, clypeus ochre, without Tibia 1 in 5 females: 8.7±10.8 (xÅ ϭ 9.6). Tip marks; sternum pale ochre with small brown of palpal tarsus as in ®g. 352. Epigynum as mark posteriorly. Distance PME-PME 0.265; in ®g. 347, ochre with distinctive light brown diameter PME 0.120; distance PME-ALE pattern; with worm-shaped ``knob'' (arrows 0.040; diameter AME 0.080. Chelicerae as in in ®gs. 347, 348). Dorsal view as in ®g. 348, P. jinwum (cf. ®g. 339), but with additional ventral cleared view as in ®g. 349. indistinct hump between proximal apophy- DISTRIBUTION: Known from several local- ses. Palps in general as in P. jinwum (cf. ®gs. ities in northern Queensland (map 17). 342, 343), only procursus distinctively dif- MATERIAL EXAMINED: AUSTRALIA: ferent (®gs. 354, 355). Legs light brown, dis- Queensland: E of Claudie Scrub, Iron Range: tal ends of femora and tibiae lighter; without Male holotype above, with 5( 3& 1 juvenile dark rings; without spines, without curved (QMB S49779); Portland Roads (12Њ35ЈS, and vertical hairs; retrolateral trichobothrium 143Њ23ЈE), May 31±June 4, 1948 (Archbold of tibia 1 at 4%; tarsus 2 distally with ϳ9 Exped.), 1& assigned tentatively, in AMNH; fairly distinct pseudosegments, proximally Wenlock River Road Xing Moreton pseudosegments not visible in dissecting mi- (12Њ03ЈS, 141Њ56ЈE), Sept. 11, 1985 (M. croscope. Opisthosoma shape as in P. jin- Bennie), 2& assigned tentatively, (QMB wum (cf. ®g. 345), with very similar brown S49777); Prince of Wales Island (10Њ41ЈS, pattern; four epiandrous spigots clearly visi- 142Њ09ЈE), July 3, 1976 (E. Cameron), 1( ble in dissecting microscope. (QMB S50230); same locality, Feb. 11, 1975 FEMALE: In general very similar to male. (E. Cameron), 1& (QMB S50219); Horn Is- Tibia 1 in 2 females: 12.7, 14.3. Epigynum land, Torres Strait (10Њ37ЈS, 142Њ17'), Dec. as in ®g. 356, ochre with distinctive light 2±8, 1986 (J. Gallon), open forest, 1( 1& brown arch in front; worm-shaped ``knob'' (QMB S12358); same locality, July 23, 1975 not visible in ventral view. Ventral cleared (H. Heatwole, E. Cameron), 2& (QMB view as in ®g. 357 (arrow points to small S50220); Normandy Stn (15Њ23ЈS, 144Њ52ЈE), knob). Sept. 13±17, 1984 (C. Fearnley), sandstone DISTRIBUTION: Known only from type lo- cliffs, 1( (QMB S49776). cality near Cairns, Queensland (map 17). MATERIAL EXAMINED: AUSTRALIA: Pholcus dungara, new species Queensland: Barron Gorge: Male holotype Figures 354±357 above, with 1( 3& (QMB S49782).

TYPE: Male holotype from Barron Gorge Pholcus koah, new species (16Њ50ЈS, 145Њ38ЈE), Queensland, Australia; Figures 358±362 Jan. 1981 (R. R. Jackson), in QMB (S34686). TYPE: Male holotype from Koah Road ETYMOLOGY: Named for the Irukandji (also (16Њ49ЈS, 145Њ31ЈE), Queensland, Australia; called Dungara), an aboriginal tribe from the Apr. 2, 1972 (N. Clyde Coleman), in QMB Cairns area, northeastern Queensland. The (S34690). species name is a noun in apposition. ETYMOLOGY: Named for the type locality. DIAGNOSIS: Very similar to P. jinwum, dis- The species name is a noun in apposition. tinguished only by the shape of the procursus DIAGNOSIS: Distinguished from the similar (compare ®gs. 343 and 355) and by the dark P. dungara and jinwum by the shape of the pattern on the epigynum (compare ®gs. 347 procursus (simple tip with two distinctive and 356). pointed terminal apophyses; ®gs. 359, 360) 114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 344±349. Pholcus jinwum. 344. Procursus, prolateral view. 345, 346. Male opisthosoma, dorsal (345) and ventral (346) views. 347±349. Epigynum, ventral (347), dorsal (348), and cleared ventral (349) views; arrows: worm-shaped ``knob''. Scale lines: 1 mm (345, 346), 0.3 mm (344, 347±349). and from P. jinwum also by the dark pattern (cf. ®gs. 342, 343); bulb almost identical (®g. on the epigynum (compare ®gs. 347 and 358); ventral femur apophysis slightly more 361). proximal; procursus distinctively different MALE (holotype): Total length 6.6, cara- (®gs. 359, 360). Legs light brown, distal ends pace width ϳ1.9 (deformed). Leg 1: 46.2 of femora and tibiae lighter; without dark (11.3 ϩ 0.9 ϩ 11.4 ϩ 19.7 ϩ 2.9), tibia 2: rings; without spines, without curved and 8.1, tibia 3 missing, tibia 4: 7.7; tibia 1 l/d: vertical hairs; retrolateral trichobothrium of 59. Habitus and prosoma shape as in P. jin- tibia 1 at 3.5%; tarsus 1 with many pseudo- wum (cf. ®gs. 339, 340). Brown mark on car- segments, but only ϳ13 distal ones distinct apace similar to P. jinwum (cf. ®g. 340), but in dissecting microscope. Opisthosoma shape without connection to ocular area; ocular as in P. jinwum (cf. ®gs. 345, 346), with very area brown, clypeus ochre, without marks; similar brown pattern. sternum light brown with ochre-yellow mar- FEMALE: In general very similar to male. gins. Distance PME-PME 0.280; diameter Tibia 1 in 4 females: 9.3, 9.5, 10.0, 10.0. PME 0.160; distance PME-ALE 0.040; di- Epigynum as in ®g. 361; ochre with light ameter AME 0.135. Chelicerae similar to P. brown arch in front, only tip of knob visible jinwum (cf. ®g. 339), but proximal apophy- in ventral view. Dorsal view as in ®g. 362 ses closer together, more pointed and direct- (arrows point to worm-shaped ``knob''). ed upward. Palps in general as in P. jinwum DISTRIBUTION: Known from three localities 2001 HUBER: PHOLCIDS OF AUSTRALIA 115

Figs. 350±353. Pholcus jinwum. 350. Male gonopore, with ®ve epiandrous spigots. 351. Pseudo- segmentation on male tarsus 2. 352. Tip of female palp. 353. Male ALS, with several piriform gland spigots. Scale lines: 60 ␮m (350, 352), 30 ␮m (351, 353). 116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

(15Њ41ЈS, 145Њ20ЈE), May 14, 1973 (V. E. Davies), 1& assigned tentatively (QMB S49775); Mt. Garnet (17Њ41ЈS, 145Њ07ЈE), Feb. 24, 1972 (N. Clyde Coleman), 1( (QMB S49773); Heron Island (23Њ27ЈS, 151Њ55ЈE), July 31, 1976 (T. Newcomb), 2& assigned tentatively (QMB S49780).

Pholcus tagoman, new species Map 17. Distribution of the genus Pholcus in Figures 363±377 Australia (except P. phalangioides). ? ϭ female specimens assigned tentatively. TYPE: Male holotype from Ningbing Cave, cave KNI-19 (15Њ18ЈS, 128Њ37ЈE), Western Australia; May 10, 1994 (R. D. Brooks), in in the Cairns area, northeastern Queensland, WAM (99/2062). and possibly from Heron Island, southeastern ETYMOLOGY: Named for the Tagoman, an Queensland (map 17). aboriginal tribe in the Daly River area in MATERIAL EXAMINED: AUSTRALIA: Northern Territory. The species name is a Queensland: Koah Road: Male holotype noun in apposition. above, with 2& (QMB S49774); Amos Bay DIAGNOSIS: Easily distinguished from Aus-

Figs. 354±357. Pholcus dungara. 354, 355. Left procursus, prolateral (354) and retrolateral (355) views. 356, 357. Epigynum, ventral (356) and cleared ventral (357) views; arrow: worm-shaped ``knob''. Scale lines: 0.3 mm. 2001 HUBER: PHOLCIDS OF AUSTRALIA 117

Figs. 358±362. Pholcus koah. 358. Left genital bulb, prolateral view; ``a'' ϭ appendix; ``e'' ϭ embolus; ``u'' ϭ uncus; ``c'' ϭ connecting piece between bulb and cymbium. 359, 360. Left procursus, prolateral (359) and retrolateral (360) views. 361, 362. Epigynum, ventral (361) and dorsal (362) views; arrows: worm-shaped ``knob''. Scale lines: 0.3 mm. tralian congeners by the absence of AME; Kritscher (1957) (possibly correctly) as Sper- from other described species also by the mophora longiventris Simon might be con- shapes of the bulbal apophyses and the pro- speci®c with the Seychellian specimens. Fi- cursus (®gs. 365, 366, 371, 372). A very nally, a single female specimen from Christ- similar species occurs on the Seychelles (M. mas Island, Phosphate Hill (10Њ26ЈS, Saaristo, unpubl. MS). It differs by having a 105Њ42ЈE) in the WAM (99/1512) might be broader uncus and an appendix ending in two conspeci®c with any of these species. tips. Two specimens from Malaysia (WAM MALE (holotype): Total length 3.8, cara- 99/1494±5) and the male identi®ed by pace width 1.0. Leg 1: 37.3 (9.2 ϩ 0.5 ϩ 9.5 118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

ϩ 16.8 ϩ 1.3), tibia 2: 5.6, tibia 3: 3.6, tibia (WAM 99/2054±61); Ningbing Cave, cave 4: 5.3; tibia 1 l/d: 101. Habitus and prosoma KNI-27 (15Њ17ЈS, 128Њ41ЈE), May 16, 1994 shape as in ®gs. 363, 364, and 367. Carapace (R. D. Brooks), 1& (WAM 99/2066); Jere- ochre, with gray-brown median stripe; ocular miah Hills, cave KJ8 (15Њ26ЈS, 128Њ44ЈE), area and clypeus without darker marks; ster- May 17, 1994 (W. F. Humphreys), 1( 1& num brown, shape as in ®g. 368. Distance (WAM 99/2068±9); same locality, May 4, PME-PME 0.215; diameter PME 0.095; dis- 1994 (B. Vine), 1& (WAM 99/2046); Point tance PME-ALE 0.025; AME absent. Che- Spring Nature Reserve (15Њ25ЈS, 128Њ53ЈE), licerae as in ®g. 369, with pair of black May 28±June 11, 1999 (M. Gray, G. Milled- apophyses distally (®gs. 370, 376) and pair ge, H. Smith), rainforest patch, 1( 1& (AMS of unsclerotized rounded apophyses proxi- KS56184±5); same data, under log, 1( 1& molaterally. Palps as in ®gs. 365 and 366, (AMS KS56186). Northern Territory: Cahills procursus and bulb as in ®gs. 371 and 372. Crossing (12Њ25ЈS, 132Њ58ЈE), May 29, 1992 Palpal tarsal organ capsulate (®g. 377), on (M. S. Harvey, J. M. Waldock), 1( (WAM conical projection of cymbium (®g. 366). 99/1513); Cutta Cutta Cave, cave K-1 Legs ochre-brown, with barely visible darker (14Њ35ЈS, 132Њ25ЈE), June 25, 1994 (B. Jones, rings on femora (subdistally) and tibiae W. Binks, R. D. Brooks), 2( 2& 6 juveniles (proximally, subdistally); without spines, (WAM 99/2035±44); Katherine, Cutta Cutta without curved and vertical hairs; retrolateral Guy Cave, 1988 (E. Holland), 1( (AMS trichobothrium of tibia 1 at 2%; tarsus 1 with KS22416); Kemp Airstrip (12Њ35ЈS, Ͼ13 pseudosegments (proximally they are 131Њ20ЈE), rainforest, Nov. 15±16, 1979 (R. dif®cult to count, distally they are quite dis- Raven), 1( (QMB S49805); Radon Creek tinct; ®g. 375). Opisthosoma shape as in ®g. (12Њ45ЈS, 132Њ53ЈE), rainforest, July 14±16, 363; gray, with blackish spots except ven- 1979 (G. Monteith, D. Cook), 1& 1 juvenile trally; genital plate not distinct. ALS with (QMB S49806). Queensland: Camooweal several piriform gland spigots. area, Great Nowranie Cave C-6 (ϳ20Њ05ЈS, VARIATION: Tibia 1 in 8 males: 6.8±9.7 (xÅ 138Њ10ЈE), 1996 (collector unknown), 1& ϭ 8.2). Opisthosoma in some males without (WAM 99/2106), assigned tentatively; same dark spots. locality, Oct. 30, 1993 (S. Eberhard), 1& FEMALE: In general very similar to male. (AMS KS40772), assigned tentatively. Tibia 1 in 18 females: 5.5±8.5 (xÅ ϭ 7.5). Epi- gynum as in ®g. 373; gray as opisthosoma PANJANGE DEELEMAN-REINHOLD AND with darker structures shining through, only DEELEMAN, 1983 knob brown to black; dorsal view as in ®g. REMARKS: I have seen only two of the six 374, apparently with divided pore plate. (Us- described species of the genus (P. mirabilis ing the light microscope I could not see and P. sedgwicki), and can therefore only add whether both plates are actually pore plates.) a few details to what has been said before by DISTRIBUTION: Apparently widely distribut- Deeleman-Reinhold and Deeleman (1983), ed in northern Northern Territory, extending Deeleman-Reinhold (1986b), and Deeleman- into northeastern Western Australia and pos- Reinhold and Platnick (1986). sibly into northwestern Queensland (map 17). Several characters support the inclusion of MATERIAL EXAMINED: AUSTRALIA: West- Panjange in the Pholcus group of genera ern Australia: Ningbing Cave, cave KNI-19: sensu Huber (1995). The ALS are provided Male holotype above, with 1& (WAM 99/ with several piriform gland spigots (®g. 389), 2063); same data but May 7±June 21, 1994 the tarsal organ is capsulate (®g. 388), and (R. D. Brooks, B. Vine, W. Binks, K. Fear, C. the male chelicerae are provided with prox- Yeates, P. Fox, W. F. Humphreys), 1( 14& imal lateral apophyses (®g. 380). The knob- several juveniles (WAM 99/2048±53, 2064, shaped apophysis at the tip of the epigynum 2067, 2070±87, 3196); Ningbing Cave, cave (®gs. 382, 383, 391) is here considered a ho- KNI-9 (15Њ17ЈS, 128Њ37ЈE), May 10, 1994 (R. mologue of a similar structure in Pholcus D. Brooks), 1& (WAM 99/ 2065); Ningbing and a few related genera, but this may be Cave, cave KNI-29 (15Њ16ЈS, 128Њ39ЈE), May wrong. In Pholcus phalangioides and P. op- 9, 1994 (R. D. Brooks), 1( 2& 5 juveniles ilionoides, this structure is grasped by the 2001 HUBER: PHOLCIDS OF AUSTRALIA 119

Figs. 363±366. Pholcus tagoman, male. 363. Habitus. 364. Prosoma, frontal view. 365, 366. Left palp, prolateral (365) and retrolateral (366) views. Scale lines: 1 mm (363), 0.4 mm (364±366). male frontal cheliceral apophyses during Panjange mirabilis Deeleman-Reinhold, copulation (Uhl et al., 1995; Huber 1995); 1986 some Panjange species have these frontal Figures 378±391 apophyses, but the species redescribed herein lacks the apophyses but nevertheless has the Panjange mirabilis Deeleman-Reinhold, 1986b: female knob-shaped apophysis. 220±222, ®gs. 45, 52±58, 59g. 120 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 367±374. Pholcus tagoman. 367. Male prosoma, dorsal view. 368. Male sternum, ventral view. 369. Male chelicerae, frontal view. 370. Distal apophysis on male chelicera. 371. Left procursus, dorsal view. 372. Left genital bulb, retrolateral view; ``a'' ϭ appendix; ``e'' ϭ embolus; ``u'' ϭ uncus; ``c'' ϭ connecting piece between bulb and cymbium. 373, 374. Epigynum, ventral (373) and dorsal (374) views. Scale lines: 0.3 mm (367±369, 371±374), 0.05 mm (370).

TYPE: Male holotype (QMB S883; right DIAGNOSIS: Distinguished from known palp and chelicerae missing) and female par- congeners by the shapes of the bulbal and atype (QMB S884) from Gordon Creek, Iron cymbial apophyses (®gs. 385, 386) (compare Range (12Њ43ЈS, 143Њ19ЈE), Queensland, ®gures of congeners in Deeleman-Reinhold Australia; June 24±30, 1976 (R. Raven, V. E. and Deeleman [1983] and Deeleman-Rein- Davies), mesophyll vine forest; examined. hold and Platnick [1986]). 2001 HUBER: PHOLCIDS OF AUSTRALIA 121

Figs. 375±377. Pholcus tagoman, male. 375. Pseudosegmentation on tarsus 1. 376. Tip of distal cheliceral apophysis. 377. Palpal tarsal organ. Scale lines: 30 ␮m (375), 15 ␮m (376, 377).

MALE (type locality, QMB S49788): Total 379), and sternum with pale ochre-yellow length 3.2, carapace width 0.93; all legs pattern. Eye triads elevated, with sclerotized loose or missing. Habitus and prosoma shape spines (®gs. 378±380); distance PME-PME as in ®gs. 378±381. Entire prosoma whitish, 0.320; diameter PME 0.120; distance PME- only carapace with light brown pattern (®g. ALE 0.030; AME absent. Sternum as in ®g. 122 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 378±384. Panjange mirabilis. 378. Habitus, male. 379±381. Male prosoma, dorsal, frontal, and ventral views. 382, 383. Epigynum, ventral and lateral views. 384. Epigynum, dorsal view. Scale lines: 1 mm (378), 0.3 mm (379±384).

381. Chelicerae whitish, with pair of pointed and extremely long apophysis that ends in apophyses proximally, otherwise unmodi®ed clawlike tip. Legs whitish, only patella area (®g. 380). Palps as in ®gs. 385 and 386, cym- and tibia±metatarsus joints darker; most hairs bium with complex procursus and simple missing. Opisthosoma shape as in ®g. 378, dorsal elongation that bears the tarsal organ whitish. at its tip; bulb with simple tubular embolus VARIATION: Carapace width in holotype: 2001 HUBER: PHOLCIDS OF AUSTRALIA 123

Figs. 385, 386. Panjange mirabilis, left male palp, prolateral (385) and retrolateral (386) views; ``e'' ϭ embolus; ``a'' ϭ bulbal apophysis; ``de'' ϭ dorsal elongation of cymbium; ``p'' ϭ procursus. Scale line: 0.5 mm.

0.84; tibia 2: 4.1, tibia 3: 2.5; the holotype mostly brown; coxae brown; opisthosoma has several dark spots on the opisthosoma. with large brown and blackish spots. Tibia 1 FEMALE: In general very similar to male, in 5 females: 5.7±6.3 (xÅ ϭ 6.0). Tarsal pseu- but eyes not elevated and closer together. Fe- dosegments dif®cult to see in dissecting mi- males from West Claudie Range with very croscope, but distinct in SEM (®g. 387). Tar- distinct brown pattern on carapace; sternum sal organ as in ®g. 388. Epigynum as in ®gs. 124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 387±391. Panjange mirabilis, female. 387. Pseudosegmentation of tarsus 1 near tip. 388. Palpal tarsal organ. 389. ALS, showing several piriform gland spigots. 390. ALS (in front) and PMS (in back). 391. Epigynal scape, ventral view. Scale lines: 60 ␮m (391), 30 ␮m (387, 390), 15 ␮m (389), 5 ␮m (388).

382, 383, and 391; dorsal view as in ®g. 384. Monteith, D. Cook), rainforest, 50 m elev., Several piriform gland spigots on ALS (®gs. 3& 1 juvenile (QMB S34664). 389, 390). DISTRIBUTION: Adults have only been col- SPERMOPHORA HENTZ, 1841 AND lected at Iron Range, northern Queensland. BELISANA THORELL, 1898 The QMB has two juveniles from Lockerbie REMARKS: Spermophora is probably the and 3 km E Lockerbie (10Њ48ЈS, 142Њ27± most chaotic genus within pholcids, and I 28ЈE) that might be conspeci®c (QMB have seen too few of the 69 species currently S4786±7) (map 19). included in Spermophora and its possible MATERIAL EXAMINED: AUSTRALIA: closest relatives (Spermophorides Wunder- Queensland: Iron Range, Gordon Creek: Male lich, Belisana Thorell, Paramicromerys Mil- holotype and female paratype above; same lot) to attempt a signi®cant improvement at data, 1( 2& several juveniles, (QMB this point. A few remarks might prove help- S49788); Iron Range, West Claudie Range ful for future workers. (12Њ45ЈS, 143Њ14ЈE), Dec. 3±10, 1985 (G. Traditionally, almost any small pholcid 2001 HUBER: PHOLCIDS OF AUSTRALIA 125

cursus like S. senoculata, but it has blackish spots like Spermophorides. The only way to solve this satisfactorily will be a worldwide revision of the genus. Unfortunately, revising just Spermophora and Spermophorides will not suf®ce, as the species described below as Belisana australis demonstrates. Belisana has been a monotypic genus since its creation by Thorell (1898), and it is with some hesitation that I assign the Australian species to this genus. I have not been able to see the type species B. taur- icornis Thorell, which Simon (1903) ®rst Maps 18, 19. Distribution of the genus Sper- mophora in Australia (map 18, top), and of the synonymized with Uthina Simon, but later genera Panjange and Belisana in Australia (map (Simon, 1909) suspected to be just another 19, bottom). Note that Belisana australis is also Spermophora. Therefore, I follow mainly the recorded from Ambon, not shown on this map. opinion of Deeleman-Reinhold (1986a: 47), who reestablished Belisana based on a col- lection of 28 morphospecies ranging from Sri with six eyes has been assigned to Spermo- Lanka to Queensland, and who kindly sent phora. So far, the New World species have me some of the specimens identi®ed by her been either removed to Metagonia or Anop- as Belisana. Deeleman-Reinhold (1986a) sicus, or synonymized with the synanthropic characterized the genitalic structure of Beli- S. senoculata (Simon, 1893; Gertsch, 1939, sana as ``somewhat similar to that in Sper- 1971, 1977, 1982; Schmidt, 1971; Yaginu- mophora'', and she (1986b: 223) used the ma, 1974; Brignoli, 1983; Huber, 2000). eye distance to separate the two genera (she However, the remaining Old World species indicated the distance between triads as be- seem to be a highly heterogenous group of ing 1±2 d in Spermophora, 3 d or more in rather small, six-eyed pholcids. Simon (1893: Belisana; this is a typing error, as the oppo- 471) noted that the genus ``n'est pas treÁs site is the case). Thus, Belisana seems to be homogeÁne'', and he distinguished two something like a pale Spermophorides, or groups: one group (including the type species better, Spermophorides a dark Belisana! S. senoculata) consisting of pale species with Again, the conclusion is that only a deep re- eye triads far apart, the other group charac- vision will clarify the issue, and this will terized by black patterns on prosoma and op- have to include Belisana as well. isthosoma and eye triads closer together. Several East-Asian Spermophora might Wunderlich (1992) formalized this distinc- actually belong to Belisana (if Deeleman's tion by creating the new genus Spermophor- interpretation proves right). For example, il- ides for the second group, but unfortunately lustrations of the genitalia of the Chinese S. he treated representatives of the Canary Is- anhuiensis Xu and Wang, of S. rollofoliolata lands almost exclusively, ignoring even the Wang, and of the Japanese S. akebona Ko- Mediterranean species. These species (S. matsu suggest a closer af®nity to Belisana huberti, mediterranea, mammata, petraea, than to Spermophora. simoni, valentiana, all described by Senglet) Some Spermophora species are too obvi- are obviously close to the Canary Island spe- ously misplaced to present a serious problem. cies, but they formally remain in Spermo- For example, judging from the existing illus- phora. More serious than that, the characters trations, S. elongata Yin and Wang and S. separating the two groups seem to work for faveauxi Lawrence are very probably Phol- Canary Island (and Mediterranean) species cus (the latter is close to, or is a synonym of versus S. senoculata, but may not work in a Pholcus circularis Kraus); S. baso Roewer worldwide perspective. For example, the new from Sumatra and S. tessellata Simon from species S. yao described below has the eye Malaysia are close relatives and certainly not triads far apart and a ventral ¯ap on the pro- Spermophora (types examined), but they 126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 probably represent a new genus; S. dubia tally and pair of rounded light apophyses Kulczynski from New Guinea is a relatively proximolaterally. Palps as in ®gs. 396 and large spider with long opisthosoma and an 397, with complex procursus; bulb with epigynum that is almost indistinguishable semitransparent projection (embolus?) and from that of Panjange mirabilis (cf. ®gs. sclerotized serrate apophysis with distal 382, 383), but the slightly damaged, opistho- hook. Legs ochre-yellow, without rings; soma of the female holotype (examined) was without spines, without curved and vertical apparently not pointed dorsoposteriorly as in hairs; retrolateral trichobothrium of tibia 1 at typical Panjange; and S. longiventris from 15%; tarsus 1 with ϳ10 fairly distinct pseu- Sumatra was described from a juvenile, and dosegments. Opisthosoma globular (®g. is probably a six-eyed Pholcus, close to P. 392); pale ochre gray, with some indistinct tagoman. large spots dorsally. ALS with several piri- At least some Spermophora species from form gland spigots, as in female. Madagascar seem to be misplaced and might VARIATION: Tibia 1 in 6 males: 1.05±1.23 end up in new genera (e.g., S. andrei, im- (xÅ ϭ 1.12). pudica), while some others (S. ankaranae, FEMALE: In general very similar to male. combesi, imerinensis, megaceros) resemble Tibia 1 in 18 females: 0.87±1.13 (xÅ ϭ 0.98). Paramicromerys in genitalic structure, a Tarsal pseudosegments as in ®g. 402. Palpal monotypic genus endemic in Madagascar. tarsal organ and tip as in ®gs. 403 and 405. Millot (1946: 146) interpreted Paramicrom- Epigynum as in ®gs. 399 and 401; frontally erys as ``faisant transition entre les Sper- brown plate with pair of pockets, posteriorly mophora et les Micromerys veÂritables'', add- dark ridges, and short, transparent scape (ar- ing yet another genus to the task awaiting the row in ®g. 401) with pocket (``p'' in ®g. 399) future revisor of Spermophora. between epigynum and spinnerets (scape hardly visible in dissecting microscope); dor- Spermophora yao, new species sal view as in ®g. 400. Several piriform Figures 392±405 gland spigots on ALS (®g. 404). DISTRIBUTION: Known only from Iron TYPE: Male holotype from Gordon Creek, Range, northern Queensland (map 18). Lower Lamond Hill, Iron Range (12Њ43ЈS, MATERIAL EXAMINED: AUSTRALIA: 143Њ19ЈE), Queensland, Australia; June 15, Queensland: Iron Range, Gordon Creek: Male 1976 (R. Raven, V. E. Davies), litter, in QMB holotype above, with 4( 12& several juve- (S34666). niles (QMB S50172); Iron Range, West ETYMOLOGY: Yao is an alternative name Claudie Range (12Њ45ЈS, 143Њ14ЈE), sieved for the Pakadji, an aboriginal tribe in north- litter in rainforest, 50 m elev., Dec. 4, 1985 eastern Queensland. The species name is a (G. Monteith), 2( 10& several juveniles noun in apposition. (QMB S33856). DIAGNOSIS: Distinguished from congeners by the shape of procursus and bulbal apoph- Spermophora paluma, new species yses (®gs. 396, 397), and by the shape of the Figures 406±415 epigynum (®g. 399). MALE (holotype): Total length 1.1, cara- TYPE: Male holotype from Paluma SF pace width 0.55. Leg 1: 4.34 (1.13 ϩ 0.19 ϩ (19Њ01ЈS, 146Њ13ЈE), Queensland, Australia; 1.15 ϩ 1.26 ϩ 0.61), tibia 2: 0.74, tibia 3: Sept. 2, 1988 (R. Raven, T.C., J. Gallon), in 0.61, tibia 4: 1.10; tibia 1 l/d: 18. Habitus QMB (S14092). and prosoma shape as in ®gs. 392±395. Car- ETYMOLOGY: Named for the type locality. apace and sternum ochre to light brown with The species name is a noun in apposition. blackish speckles, ocular area and clypeus DIAGNOSIS: Distinguished from known without speckles (®gs. 393±395). Eye triads congeners by the shape of procursus and bul- hardly elevated; distance PME-PME 0.215; bal apophyses (®gs. 408±410), and by the diameter PME 0.050; distance PME-ALE shape of the epigynum (®g. 412). The AMS 0.010; AME absent. Chelicerae as in ®g. 398, has a very close undescribed relative from with pair of simple pointed apophyses dis- Lord Howe Island (31Њ33ЈS, 159Њ05ЈE) 2001 HUBER: PHOLCIDS OF AUSTRALIA 127

Figs. 392±396. Spermophora yao, male. 392. Habitus. 393. Prosoma, frontal view. 394. Sternum, ventral view. 395. Prosoma, dorsal view. 396. Left palp, retrolateral view; ``e'' ϭ putative embolus; ``h'' ϭ bulbal apophysis with hooked tip; ``f'' ϭ ventral ¯ap. Scale lines: 0.3 mm.

(KS56205, 56217) that is signi®cantly larger pace width ϳ0.56 (deformed). Leg 1: 17.2 (total length 2±2.1; carapace width 0.8; tibia (4.3 ϩ 0.3 ϩ 4.1 ϩ 7.3 ϩ 1.2), tibia 2: 2.6, 1: 4.5±4.6) and has only one tip on the pu- tibia 3: 1.6, tibia 4: 2.7; tibia 1 l/d: 68. Pro- tative embolus (instead of two as in ®gs. 408, soma shape as in ®gs. 406 and 407; entire 409); chelicerae and procursus are almost prosoma pale ochre; sternum whitish. Eye identical to S. paluma. triads hardly elevated; distance PME-PME MALE (holotype): Total length 1.4, cara- 0.185; diameter PME 0.070; distance PME- 128 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 397±400. Spermophora yao. 397. Left male palp, prolateral view. 398. Male chelicerae, frontal view. 399. Epigynum with posterior pocket (``p''), ventral view. 400. Epigynum, dorsal view. Scale lines: 0.3 mm (397), 0.2 mm (398±400).

ALE 0.010; AME absent. Chelicerae as in gonopore with four epiandrous spigots (®g. ®g. 411, with pair of simple pointed apoph- 413); several piriform gland spigots on ALS yses distally and pair of rounded light apoph- (®g. 415). yses proximolaterally. Palps as in ®g. 408, VARIATION: Tibia 1 in 10 males: 3.5±4.1 with complex procursus; bulb with two sim- (xÅ ϭ 3.8). Some males with large blackish ple projections, one of them (putative em- spots on opisthosoma. bolus) ending in two black tips (®gs. 408 and FEMALE: In general very similar to male. 409); procursus as in ®gs. 408±410. Legs Epigynum as in ®g. 412; extremely pale, pale ochre-yellow, patella area slightly dark- with pair of pockets in frontal plate and un- er, ventroproximal margin dark brown; with- paired pocket on short, transparent posterior out spines, without curved and vertical hairs; scape between epigynum and spinnerets retrolateral trichobothrium of tibia 1 at 3%; (``p'' in ®g. 412). tarsus 1 distally with ϳ10 fairly distinct DISTRIBUTION: Known from several local- pseudosegments, proximally possibly more. ities in northeastern Queensland (map 18). Tarsal organ as in ®g. 414. Opisthosoma pale MATERIAL EXAMINED: AUSTRALIA: gray, shape similar to S. yao (cf. ®g. 392), Queensland: Paluma SF: Male holotype above, but slightly longer (0.93 long, 0.67 high); with 2( (QMB S14092); Wallaman Falls via 2001 HUBER: PHOLCIDS OF AUSTRALIA 129

Figs. 401±405. Spermophora yao, female. 401. Epigynum and posterior scape (arrow), ventral view. 402. Pseudosegmentation of tarsus (unidenti®ed leg). 403. Tip of palp. 404. ALS, showing several piriform gland spigots. 405. Palpal tarsal organ. Scale lines: 40 ␮m (401, 402), 5 ␮m (403±405).

Ingham (18Њ36ЈS, 145Њ48ЈE), rainforest, 500 teith), 3( 3 juveniles (QMB S50241); 3 km m elev., Oct. 1, 1980 (G. Monteith), 1( 3 N of Kuranda, Kuranda State Forest juveniles (QMB S50247); Mt. Graham (ϳ16Њ47ЈS, 145Њ38ЈE), rainforest, 360 m (18Њ23ЈS, 145Њ52ЈE), Dec. 26±30, 1986 (S. elev., June 25±Oct. 3, 1982 (S. & J. Peck), Hamlet), rainforest, 600±700 m elev., 1( 3 1( (AMNH); 20 km SW of Mossman, Mt. juveniles (QMB S18422); Kirrama Range Lewis (ϳ16Њ35ЈS, 145Њ15ЈE), rainforest, 900 (18Њ12ЈS, 145Њ45ЈE), Douglas Creek Road, m elev., June 26±Oct. 1, 1982 (S. & J. Peck), 800 m elev., Dec. 9±12, 1986 (G. Monteith, 1( (AMNH); Fritz Creek, N of Bloom®eld G. Thompson, S. Hamlet), 1( (QMB (15Њ52ЈS, 145Њ21ЈE), rainforest foliage, Dec. S49803); Kirrama Range via Kennedy 1975 (M. Gray), 1( (AMS KS65695). (ϳ18Њ09ЈS, 145Њ43ЈE), rainforest, 500 m elev., Oct. 2, 1980 (G. Monteith), 1( 1 ju- Belisana australis, new species venile (QMB S49760); Boonjee (17Њ24ЈS, Figures 416±429 145Њ44ЈE), Apr. 3±6, 1978 (V. E. Davies, R. TYPE: Male holotype from Home Rule Raven), 2( 1 juvenile (QMB S50270); Win- (15Њ44ЈS, 145Њ18ЈE), Queensland, Australia; din Falls, via Butchers Creek (17Њ19ЈS, Oct. 28±Nov. 18, 1974 (V. E. Davies), litter, 145Њ45ЈE), rainforest, Oct. 9, 1980 (G. Mon- in QMB (S50273). 130 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 406±412. Spermophora paluma. 406, 407. Male prosoma, frontal and dorsal views. 408. Left male palp, retrolateral view. 409. Left procursus and genital bulb, prolateral view. 410. Left cymbium and procursus, dorsal view; ``f'' ϭ ventral ¯ap. 411. Male chelicerae, frontal view. 412. Epigynum, ventral view; ``p'' ϭ pocket. Scale lines: 0.3 mm (406±410), 0.1 mm (411, 412). 2001 HUBER: PHOLCIDS OF AUSTRALIA 131

Figs. 413±415. Spermophora paluma, male. 413. Gonopore, showing four epiandrous spigots. 414. Tarsal organ on tarsus 2. 415. ALS, showing several piriform gland spigots. Scale lines: 20 ␮m (413), 5 ␮m (414).

ETYMOLOGY: The species name (Latin aus- described pholcids where the male is known tralis, ``southern'') refers to the geographic by the huge apophysis on the male palpal distribution of this southernmost Belisana trochanter (®g. 419). The collection CLD has species. a male of a probably undescribed species DIAGNOSIS: Tiny six-eyed pholcid with from Thailand with a similar apophysis, but globular opisthosoma; distinguished from all with very different chelicerae. 132 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

MALE (holotype): Total length 1.2, cara- 132Њ16ЈE), rainforest, Nov. 12, 1979 (R. Ra- pace width 0.52. Leg 1: 8.65 (2.19 ϩ 0.19 ϩ ven), 3& 2 juveniles (QMB S50173); same 2.24 ϩ 3.03 ϩ 1.00), tibia 2: 1.42, tibia 3: locality, Jan. 1979 (R. Raven), 1& 2 juve- 0.87, tibia 4: 1.34; tibia 1 l/d: 48. Habitus niles (QMB S50180); West Alligator River, and prosoma shape as in ®gs. 416, 417, and ``mouth, WA1'' (12Њ12ЈS, 132Њ13ЈE), Nov. 420. Carapace light ochre, with slightly dark- 12, 1979 (R. Raven), 1( 1& 5 juveniles er bands posteriorly and around ocular area (QMB S50175); East Alligator Xing (®g. 420); ocular area and clypeus without (12Њ25ЈS, 132Њ58ЈE), sieved litter in rainfo- markings; sternum pale ochre. Ocular area rest, July 18, 1979 (G. B. Monteith), 2( 6 not elevated; distance PME-PME 0.105; di- juveniles (QMB S50179); North Point, Ka- ameter PME 0.070; distance PME-ALE palga (12Њ25ЈS, 132Њ22ЈE), rainforest, Nov. 0.020; AME absent. Chelicerae as in ®g. 423, 11, 1979 (R. Raven), 1( 1& (QMB S50171); with pair of simple pointed apophyses Radon Creek (12Њ45ЈS, 132Њ53ЈE), Nov. 14, distally and pair of rounded light apophyses 1979 (R. Raven), litter, wet rainforest, 2( 3 proximolaterally. Palps as in ®gs. 418 and juveniles (QMB S50177). INDONESIA: 419, with huge trochanter apophysis and Moluccas: Ambon, Jan. 17, 1995 (C. L. & P. small apophysis on femur retrolaterodorsally; R. Deeleman), 1( (CLD). procursus and bulb relatively simple (®gs. 421, 422). Legs light ochre, without rings; INTRODUCED SPECIES without spines, curved, and vertical hairs; Pholcids are among the most common spi- retrolateral trichobothrium of tibia 1 at 18%; ders in houses throughout the world, and tarsus 1 distally with ϳ14 fairly distinct most of the synanthropic species have been pseudosegments (®g. 427), proximally pseu- found in Australia. Synanthropics make up a dosegmentation dif®cult to see in dissecting substantial fraction of most collections. Be- microscope. Opisthosoma almost globular cause the priority of this work was to estab- (®g. 416), ochre-gray; ALS with several pir- lish the diversity of the endemic fauna, I iform gland spigots; gonopore with four have rather neglected the introduced species, epiandrous spigots in two groups. and in some cases I have even asked curators VARIATION: Tibia 1 in 5 males: 1.7±2.5 (xÅ not to send me the reliably identi®ed ones. ϭ 2.13). Nevertheless, they are all included in the key FEMALE: In general very similar to male. above, and a few additional notes might Palpal tarsus tip as in ®g. 429. Epigynum as prove useful for future workers. in ®g. 424, with distinctive scape provided For practical purposes, the nine introduced with distal pair of pockets (®g. 425); dorsal pholcid species of Australia can be divided view as in ®g. 426. Several piriform gland into two groups: the common species (the spigots on ALS (®g. 428). ®rst six below), living usually in or around DISTRIBUTION: Known from several local- houses; and the rare ones (three), found just ities in Australia (northern Northern Territo- in one or a few occasions, apparently not so ry and northern Queensland; map 19) and closely associated with humans. Following is from Ambon, Moluccas. a list of all species, sorted roughly from com- MATERIAL EXAMINED: AUSTRALIA: mon to rare, with references to published il- Queensland: Home Rule: Male holotype lustrations useful for identi®cation and to above, with 1& (epigynum lost) in same vial; their probable geographic origin. Clifton Beach (16Њ46ЈS, 145Њ40ЈE), 1971± 1972 (N. Clyde-Coleman), 1& (QMB Pholcus phalangioides (Fuesslin, 1775) S50284); West Claudie Range, Iron Range (12Њ45ЈS, 143Њ14ЈE), Dec. 3±10, 1985 (G. Illustrations: Wiehle, 1953: ®gs. 102±107; Uhl et Monteith, D. Cook), rainforest, 3( 3 juve- al., 1995: ®gs. 2, 3, 5. niles (QMB S34660); Murray Island, Torres This species is commonly seen as an orig- Strait (9Њ56ЈS, 144Њ04ЈE), Aug. 1974 (H. inally European species, but the illustrations Heatwole, E. Cameron), 1( 1 juvenile of Chinese and Japanese Pholcus species (QMB S50231). Northern Territory: West (e.g., Zhu et al., 1986; Zhu and Gong, 1991; Alligator River, ``mouth, WA2'' (12Њ11ЈS, Irie, 1997; Song et al., 1999) suggest that the 2001 HUBER: PHOLCIDS OF AUSTRALIA 133

Figs. 416±419. Belisana australis, male. 416. Habitus. 417. Prosoma, frontal view. 418, 419. Left palp, prolateral (418) and retrolateral (419) views. Scale lines: 0.5 mm (416), 0.3 mm (417±419).

closest relatives might as well be East Asian. narrow uncus (®gs. 342, 358), and their epi- Whatever the origin, P. phalangioides is gynum is roundish and weakly sclerotized quite different from the eight-eyed Australian (®gs. 347, 356, 361) rather than triangular Pholcus species and is thus easily distin- and more heavily sclerotized, as in P. phal- guished; that is, Australian species have a angioides. 134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Figs. 420±426. Belisana australis. 420. Male prosoma, dorsal view. 421. Left genital bulb, dorsal view. 422. Left procursus, prolateral view. 423. Male chelicerae, frontal view. 424. Epigynum, ventral view. 425. Epigynal scape, ventral view. 426. Epigynum, dorsal view. Scale lines: 0.3 mm (420, 424), 0.1 mm (421±423, 426).

Smeringopus pallidus (Blackwall, 1858) in Australia (see below) suggests that some Figures 431, 434±436 old records of S. pallidus might actually re- sult from misidenti®cations. However, in Further illustrations: Millot, 1941: ®gs. 7A±I (as S. elongatus); Kraus, 1957: ®gs. 1±6; Saaristo, Australia, S. pallidus is incomparably more 1978: ®gs. 23±26, 31±38. common, and, in contrast to S. natalensis, is closely associated with humans. It is eas- Smeringopus is an African (perhaps only ily distinguished from S. natalensis and sub-Saharan) genus. Only S. pallidus has other congeners by the bulbal apophyses previously been known to have been intro- (®gs. 434, 435), the tip of the procursus duced to other continents. The discovery of (®g. 431), and the shape of the epigynum the super®cially very similar S. natalensis (®g. 436). 2001 HUBER: PHOLCIDS OF AUSTRALIA 135

Figs. 427±429. Belisana australis. 427. Pseudosegmentation of male tarsus 1. 428. Female ALS, showing several piriform gland spigots. 429. Tip of female palp, with tarsal organ. Scale lines: 20 ␮m (427), 10 ␮m (428, 429).

Artema atlanta Walckenaer, 1837 This possibly biggest of all pholcid species is easily distinguished from other Australian Illustrations: Millot, 1941: ®gs. 1A±I (as A. maur- iciana), 1946: ®g. 1 (as A. mauriciana); Brig- pholcids by its large size. Both the procursus noli, 1981: ®gs. 1±7. (Millot, 1941: ®gs. 1A, C) and the epigynum 136 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

(Millot, 1946: ®g. 1) are unique. It probably interesting to note that even there it is a originated from the Middle East. Its (poorly ``semi-domesticated species'' (Lawrence, known) congeners are from Iran, Afghani- 1967: 299), ``common in buildings . . . in stan, Turkmenistan, and Tadzhikistan. various rooms and outhouses'' (Lawrence, 1947: 15). Given this apparent preadaptation Crossopriza lyoni (Blackwall, 1867) to live with humans, it seems surprising that Illustrations: Millot, 1946: ®gs. 29A, B, 30B, this species has not been found on other con- 31A, B (as C. francoisi and C. stridulans). tinents before. However, as mentioned above, some records of S. pallidus might actually This species probably originated form result from misidenti®cations of S. natalen- northern Africa, the Mediterranean region, or sis. In Australia, S. natalensis has been col- the Middle East. It is most easily distin- lected at various localities, some of them guished by the two pairs of distinctive densely populated urban areas (M. Gray, per- apophyses on the male chelicerae (Millot, sonal commun.), but apparently never in 1946: ®g. 29A) and by the shape of the epi- houses. It is easily distinguished from S. pal- gynum (Millot, 1946: ®g. 31B). lidus as shown in ®gs. 430±437. MATERIAL EXAMINED: SOUTH AFRICA: Physocyclus globosus (Taczanowski, 1874) Zululand, Kosi Bay Reserve, Kosi Lakes Illustrations: Brignoli, 1981: ®gs. 14±18, 21±24; (27Њ00ЈS, 32Њ24ЈE), June 5±15, 1995 (E. Huber and Eberhard, 1997: ®gs. 1, 2, 7A, B, 9. & R. Kyle, R. A. Cooper), 1( 3& (WAM 99/1482±7). AUSTRALIA: Western Austra- Physocyclus has a much more restricted lia: Tuart Hill near Perth (31Њ53ЈS, distribution (western USA to Costa Rica) 115Њ51.5ЈE), Sept. 24, 1993±May 21, 1994 than previously thought, and P. globosus is (M. S. Harvey, J. M. Waldock), 3( 4& ϳ10 probably the only species found all over the juveniles (WAM 99/2117±2135). New South world in tropical and subtropical climates. Wales: Ashford, Ashford Cave (29Њ12ЈS, Physocyclus appears to be closely related to 151Њ00ЈE), Jan. 31, 1995 (S. Eberhard), 1( the Australian genus Trichocyclus, but P. 1 juvenile (AMS KS49265); Malabar globosus is easily distinguished from Tricho- (33Њ58ЈS, 151Њ15ЈE), June 26 and Sept. 28, cyclus species and other Australian pholcids 1966 (R. E. Mascord), 3& (AMS KS48725, by the bulbal apophysis, the shape of the pro- 56197); Penrith (33Њ45ЈS, 150Њ42ЈE), Aug. cursus, and the epigynum (Huber and Eber- 25, 1979 (A. Johnson), 1& (AMS KS65700). hard, 1997: ®gs. 1, 7). Micropholcus fauroti (Simon, 1887) Holocnemus pluchei (Scopoli, 1763) Illustrations: Millot, 1946: ®gs. 2A, B (as Pholcus Illustrations: Brignoli, 1971: ®gs. 1±6; Timm, chavanei); Deeleman-Reinhold and Prinsen, 1976: ®gs. 1, 2; Huber, 1995: ®gs. 1A, 2A, B, 1987: ®gs. 1±9. 4A, B. Since Micropholcus is a monotypic genus, Holocnemus is closely related to (possibly and the sister group is not known, the origin a synonym of) Crossopriza, and shares the of the pantropical M. fauroti is obscure. The northern African and Mediterranean distri- species is new for Australia, and is repre- bution. The only cosmopolitan species, H. sented in the collections studied by only two pluchei, is easily distinguished from other females from Western Australia, Broome, Australian pholcids by the bulbal apophyses Cable Beach (17Њ57ЈS, 122Њ12ЈE), July 9, and the small cheliceral apophyses (Huber, 1981 (D. Hirst), ``on rocks, base of which 1995: ®gs. 2B, 4A). are at high tide mark'', in SAM (N1999/862± 3). Males of this species are easily distin- Smeringopus natalensis Lawrence, 1947 guished by their long dorsal hinged process Figures 430, 432, 433, 437 on the procursus (Millot, 1946: ®g. 2A). Fe- males are much more dif®cult to identify, but Further illustrations: Lawrence, 1967: ®gs. 3±5. have a characteristic triangular structure in This species has previously been known the internal genitalia (Deeleman-Reinhold only from eastern South Africa (Natal). It is and Prinsen, 1987: ®g. 7). 2001 HUBER: PHOLCIDS OF AUSTRALIA 137

Figs. 430±437. Smeringopus natalensis (430, 432, 433, 437), S. pallidus (431, 434±436). 430, 431. Left procursi, retrolateral views. 432, 434. Left genital bulbs, retrolateral views. 433, 435. Left genital bulbs, prolateral views. 436, 437. Epigyna, ventral views. Scale lines: 0.3 mm (430±435), 0.5 mm (436, 437).

Modisimus culicinus (Simon, 1893) short-legged representatives similar to M. Illustrations: Huber, 1997a: ®gs. 2±4, 1997b: culicinus (mostly undescribed) are diverse on ®g. 1. the Antilles and in Florida. Outside this area, M. culicinus has been recorded from various Modisimus is a New World genus, and 138 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260 countries in the Americas and from the Con- Brignoli, P. M. go, the Seychelles, and Micronesia; it is new 1971. Note sui Pholcidae d'Italia. Fragm. En- to Australia. The collections studied contain tomol. 7(2): 79±101. a male from Queensland, Rose Bay near 1980. Sur le genre Leptopholcus Simon, 1893 Bowen (20Њ00ЈS, 148Њ16ЈE), July 27±Dec. 2, (Araneae, Pholcidae). Rev. Zool. Afr. 1992 (R. Raven, P. & E. Lawless, M. Shaw), 94(3): 649±655. 1981. Studies on the Pholcidae, I. Notes on in QMB (S24958); a male from Queensland, the genera Artema and Physocyclus Mt. Molloy (16Њ44ЈS, 145Њ19ЈE), 400 m (Araneae). Bull. Am. Mus. Nat. Hist. elev., ``riparian/woodland'', 1992±1993 (S. 170(1): 90±100. Burnett), in QMB (S33178); and a female 1983. A catalogue of Araneae described be- from Papua New Guinea, Central Province, tween 1940 and 1981. Manchester: Sept. 1, 1985 (D. Court) in the CLD collec- Manchester Univ. Press. 755 pp. tion. Males of this species are easily distin- Brown, D. A., K. S. W. Campbell, and K. A. W. guished by the cuticular lobe on the clypeus Crook (Huber, 1997b: ®g. 1). 1968. The Geological Evolution of Australia and New Zealand. New York: Perga- mon. ACKNOWLEDGMENTS Deeleman-Reinhold, C. L. 1986a. Leaf-dwelling Pholcidae in Indo-Aus- I am deeply indebted to Norman Platnick tralian rain forests. Proc. IX Int. Congr. for his support throughout this study. Nu- Arachn. (Panama, 1983): 45±48. merous others assisted in uncountable ways 1986b. Studies on tropical Pholcidae. II. Re- and made my stay at the American Museum description of Micromerys gracilis of Natural History a pleasure, namely Diana Bradley and Calapnita vermiformis Si- Silva, Tam Nguyen, Vladimir Ovtsharenko, mon (Araneae, Pholcidae) and descrip- Xin-Ping Wang, and Boris Zakcharov. Spe- tion of some related new species. Mem. cial thanks to Lou Sorkin for help with all Queensland Mus. 22(2): 205±224. sorts of problems, and to Angela Klaus for 1993. Description of a new cave-dwelling her assistance with the SEM. pholcid spider from north-western Aus- I depended on the time and effort of cu- tralia, with an identi®cation key to the rators, curatorial assistants, and other indi- genera of Australian Pholcidae (Ara- neae). Rec. West. Aust. Mus. 16(3): viduals to send me material, and I want to 323±329. thank all those who answered my requests: 1995. Redescription of Holocneminus multi- Jonathan Coddington (USNM), Hieronymus guttatus Simon and description of two Dastych (ZMH), Christa Deeleman-Reinhold new species of pholcid spiders from (Netherlands), Mike Gray and Graham Mil- Australia. Beitr. Araneol. 4: 31±41. ledge (AMS), Mark Harvey and Julianne Deeleman-Reinhold, C. L., and P. R. Deeleman Waldock (WAM), David Hirst (SAM), Rob- 1983. Studies on tropical Pholcidae. I. Pan- ert Raven (QMB), and Christine Rollard jange, a new genus of Indo-Australian (MNHN). leaf- and rock-dwelling pholcid spiders Comments and suggestions by Norman (Araneae). Zool. Meded. 57(14): 121± Platnick, Mark Harvey, Michael Gray, and 130. Jonathan Coddington greatly improved the Deeleman-Reinhold, C. L., and N. I. Platnick 1986. A new Panjange from northern Borneo manuscript. The research was funded by a (Araneae, Pholcidae). J. New York En- Kalb¯eisch Research Fellowship at the tomol. Soc. 94(4): 559±561. American Museum of Natural History, New Deeleman-Reinhold, C. L., and J. D. Prinsen York. 1987. Micropholcus fauroti (Simon) n. comb., a pantropical, synanthropic spi- REFERENCES der (Araneae: Pholcidae). Entomol. Ber., Amsterdam 47(5): 73±77. Bradley, H. H. B. Farris, J. S. 1877. The Araneides of the ``Chevert'' Ex- 1988. Hennig86, version 1.5. Program and pedition. Part II. Proc. Linn. Soc. New documentation. Port Jefferson, New South Wales 2: 115±120. York. 2001 HUBER: PHOLCIDS OF AUSTRALIA 139

Gertsch, W. J. Jackson, R. R. 1939. A new genus in the Pholcidae. Am. 1992. Predator-prey interactions between Mus. Novitates 1033: 4 pp. web-invading jumping spiders and two 1971. A report on some Mexican cave spi- species of tropical web-building phol- ders. Assoc. Mex. Cave Stud. Bull. 4: cid spiders, Psilochorus sphaeroides 47±111. and Smeringopus pallidus. J. Zool. 1977. Report on cavernicole and epigean spi- (London) 227: 531±536. ders from the Yucatan Peninsula. In J. Jackson, R. R., and R. J. Rowe R. Reddell (ed.), Studies on the caves 1987. Web-invasion and araneophagy by New and cave fauna of the Yucatan Penin- Zealand and Australian pholcid spiders. sula. Assoc. Mex. Cave Stud. Bull. 6: New Zealand J. Zool. 14: 139±140. 103±131. Jackson, R. R., R. J. Rowe, and G. E. Campbell 1982. The spider genera Pholcophora and 1992. Anti-predator defenses of Psilochorus Anopsicus (Araneae, Pholcidae) in sphaeroides and Smeringopus pallidus North America, Central America and (Araneae, Pholcidae), tropical web- the West Indies. Bull. Texas Mem. building spiders. J. Zool. (London) Mus. 28: 95±144. 228: 227±232. Goloboff, P. A. Keast, A. 1993. NONA, version 1.8. Program and doc- 1981a. Distributional patterns, regional biotas, umentation. New York, New York. and adaptations in the Australian biota: 1997. Pee-Wee, version 2.8. Program and a synthesis. In A. Keast (ed.), Ecolog- documentation. New York, New York. ical biogeography of Australia: 1891± Huber, B. A. 1997. The Hague: W. Junk Publ. 1995. Copulatory mechanism in Holocnemus 1981b. Origins and relationships of the Austra- pluchei and Pholcus opilionoides, with lian biota. In A. Keast (ed.), Ecological notes on male cheliceral apophyses and biogeography of Australia: 2001±2050. The Hague: W. Junk Publ. stridulatory organs in Pholcidae (Ara- Koch, L. neae). Acta Zool. (Stockholm) 76(4): 1867. Beschreibung neuer Arachniden und 291±300. Myriapoden. Verh. Zool. Bot. Ges. 1997a. On the distinction between Modisimus Wien 17: 173±250. and Hedypsilus (Pholcidae; Araneae), 1872. Die Arachniden Australiens. Bauer und with notes on behavior and natural his- Raspe, NuÈrnberg, 1: 105±368. tory. Zool. Scr. 25(3): 233±240. Kraus, O. 1997b. Evidence for gustatorial courtship in a 1957. Araneenstudien 1. Pholcidae (Smerin- haplogyne spider (Hedypsilus culici- gopodinae, Ninetinae). Senckenb. Biol. nus: Pholcidae: Araneae). Netherlands 38(3/4): 217±243. J. Zool. 47(1): 95±98. Kritscher, E. 1997c. On American `Micromerys' and Meta- 1957. Bisher unbekannt gebliebene Araneen- gonia (Araneae, Pholcidae), with notes MaÈnnchen und -Weibchen des Wiener on natural history and genital mechan- Naturhistorischen Museums (1. Teil). ics. Zool. Scr. 25(4): 341±363. Ann. Naturhist. Mus. Wien 61: 254± 1998. On the ``valve'' in the genitalia of fe- 272. male pholcids (Pholcidae, Araneae). Lawrence, R. F. Bull. Br. Arachnol. Soc. 11(2): 41±48. 1947. A collection of Arachnida made by Dr. 2000. New World pholcid spiders (Araneae: I. TraÈgardh in Natal and Zululand Pholcidae): a revision at generic level. (1904±1905). K. Vet. Vitterh. Samh. Bull. Am. Mus. Nat. Hist. 254: 1±348. Handl. (ser. B) 5(9): 3±41. Huber, B. A., and W. G. Eberhard 1967. A new cavernicolous Pholcid spider 1997. Courtship, copulation, and genital me- from the Congo. Rev. Suisse Zool. chanics in Physocyclus globosus (Ara- 74(4): 295±300. neae, Pholcidae). Can. J. Zool. 74: Millot, M. J. 905±918. 1941. Les araigneÂes de l'Afrique occidentale Irie, T. Francaise. Sicariides et pholcides. 1997. Two new species of the genera Pholcus MeÂm. Acad. Sci. Inst. France 64: 1±30. and Spermophora (Araneae: Pholcidae) 1946. Les pholcides de Madagascar (AraneÂi- from the Kyushu, Japan. Acta Arach- des). MeÂm. Mus. Natl. Hist. Nat. (n. nol. 46(2): 133±138. ser.) 22(3): 127±158. 140 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

Nix, H. A. Pholcidae). Bull. Br. Arachnol. Soc. 1981. The environment of Terra Australis. In 10(1): 1±9. A. Keast (ed.), Ecological biogeogra- Wiehle, H. phy of Australia: 105±133. The Hague: 1953. Spinnentiere oder Arachnoidea (Ara- W. Junk Publ. neae), IX. In M. Dahl and H. Bischoff Nixon, K. C. (eds.), Die Tierwelt Deutschlands. 1992. Clados, version 1.2. Program and doc- Jena: Gustav Fischer. 150 pp. umentation. Trumansburg, New York. Wunderlich, J. 1999. Winclada, version 0.9.9 ϩ. Program 1992. Die Spinnen-Fauna der Makaronesisch- and documentation. Ithaca, New York. en Inseln. Taxonomie, OÈ kologie, Bio- Paik, K. Y. geographie und Evolution. Beitr. Ara- 1978. The Pholcidae (Araneae) of Korea. neol. 1: 1±620. Educ. J. 20(1978): 113±135. Yaginuma, T. Rack, G. 1974. The spider fauna of Japan (IV). Fac. 1961. Die entomologischen Sammlungen des Let. Rev. Otemon Gakuin Univ. 8: Zoologischen Staatsinstituts und Zool- 169±173. ogischen Museums Hamburg. Mitt. Zhu, M., and L.-S. Gong Hamburg. Zool. Mus. Inst. 59: 1±60. 1991. Four new species of the genus Pholcus Rainbow, W. J. from China. Acta Zootaxonomica Sin- 1911. A census of Australian Araneidae. Rec. ica 16(1): 18±26. Aust. Mus. 9(2): 107±319. Zhu, M., H. Tu, and J. G. Shi Saaristo, M. I. 1986. Spiders of the genus Pholcus (Araneae, 1978. Spiders (Arachnida, Araneae) from the Pholcidae) from Shanxi and Shanxi Seychelle Islands, with notes on tax- Provinces, China, with description of a onomy. Ann. Zool. Fenn. 15: 99±126. new species. J. Hebei Normal Univ. Schmidt, G. 1986(2): 118±124. 1971. Mit Bananen eingeschleppte Spinnen. Zool. Beitr. 17: 387±433. INDEX OF GENERIC AND SPECIFIC Simon, E. 1893. Histoire naturelle des araigneÂes 1(2): NAMES 256±488. Paris. 1903. Histoire naturelle des araigneÂes 2(4): Page numbers refer to the ®rst mention of 669±1080. Paris. a species. 1908. Araneae, 1re partie. In W. Michaelsen and R. Hartmeyer (eds.), Die Fauna arabana, n. sp. (Trichocyclus), 65 SuÈdwest-Australiens 1(12): 359±446. aranda, n. sp. (Trichocyclus), 75 Jena: Gustav Fischer. arawari, n. sp. (Trichocyclus), 86 Â arcoona, n. sp. (Wugigarra), 36 1909. Etude sur les archnides du Tonkin (1eÁre arnga, n. sp. (Trichocyclus), 90 partie). Bull. Sci. Fr. Belg. 42: 69±147. australis, n. sp. (Belisana), 129 Song, D., M. Zhu, and J. Chen 1999. The spiders of China. Hebei Science balladong, n. sp. (Trichocyclus), 75 and Technology Publishing House, 640 Belisana Thorell, 124 pp., 4 pls. bugai, n. sp. (Trichocyclus), 90 Thorell, T. bujundji, n. sp. (Wugigarra), 42 1898. Secondo saggio sui ragni Birmani. II. bulburin, n. sp. (Wugigarra), 33 burgul, n. sp. (Wugigarra), 49 Retitelariae et Orbitelariae. Ann. Mus. Civ. Genova 39: 271±378. daviesae Deeleman-Reinhold (Micromerys), 107 Timm, H. djauan, n. sp. (Trichocyclus), 76 1976. Die Bedeutung von Genitalstrukturen dungara, n. sp. (Pholcus), 113 fuÈr die KlaÈrung systematischer Fragen bei Zitterspinnen (Arachnida: Araneae: eberhardi, n. sp. (ugigarra), 22 Pholcidae). Entomol. Germ. 3(1/2): 69±76. gia, n. sp. (ugigarra), 39 gidil, n. sp. (Micromerys), 105 Uhl, G., B. A. Huber, and W. Rose gnalooma, n. sp. (Trichocyclus), 78 1995. Male pedipalp morphology and copu- gracilis Bradley (Micromerys), 98 latory mechanism in Pholcus phalan- grayi, n. sp. (Trichocyclus), 83 gioides (Fuesslin, 1775) (Araneae, gurran, n. sp. (Micromerys), 103 2001 HUBER: PHOLCIDS OF AUSTRALIA 141

Pholcus Walckenaer, 108 harveyi, n. sp. (Trichocyclus), 92 pustulatus Deeleman-Reinhold (Trichocyclus), 86 hirsti n. sp. (Trichocyclus), 62 raveni, n. sp. (Micromerys), 107 idi, n. sp. (Wugigarra), 48 septentrionalis Deeleman-Reinhold (Trichocyclus), 80 jiman, n. sp. (Wugigarra), 30 Spermophora Hentz, 124 jinwum, n. sp. (Pholcus), 111 sphaeroides (Koch) (Wugigarra), 20 kalamai, n. sp. (Wugigarra), 54 tagoman, n. sp. (Pholcus), 116 kaurna, n. sp. (Wugigarra), 15 tjapukai, n. sp. (Wugigarra), 13 koah, n. sp. (Pholcus), 113 Trichocyclus Simon, 56 kokata, n. sp. (Trichocyclus), 64 kurara, n. sp. (Trichocyclus), 80 undanbi, n. sp. (Wugigarra), 29 ungumi, n. sp. (Trichocyclus), 92 mamu, n. sp. (Wugigarra), 14 wanjuru, n. sp. (Wugigarra), 49 Micromerys Bradley, 95 warianga, n. sp. (Trichocyclus), 73 mirabilis Deeleman-Reinhold (Panjange), 119 watta, n. sp. (Trichocyclus), 95 muluridji, n. sp. (Wugigarra), 39 wigi, n. sp. (Micromerys), 105 wiri, n. sp. (Wugigarra), 32 nauo, n. sp. (Wugigarra), 51 worora, n. sp. (Trichocyclus), 89 nigropunctatus Simon (Trichocyclus), 70 Wugigarra, n. gen., 11 nullarbor, n. sp. (Trichocyclus), 59 wulpura, n. sp. (Wugigarra), 47 wunderlichi (Deeleman-Reinhold) (Wugigarra), 45 oborindi, n. sp. (Trichocyclus), 83 yao, n. sp. (Spermophora), 126 paluma, n. sp. (Spermophora), 126 yawai, n. sp. (Wugigarra), 24 pandima, n. sp. (Trichocyclus), 65 yidin, n. sp. (Micromerys), 102 Panjange Deeleman-Reinhold and Deeleman, 118 yirgai, n. sp. (Wugigarra), 34 142 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

APPENDIX 1 Additions to the Matrix in Huber (2000) 2001 HUBER: PHOLCIDS OF AUSTRALIA 143

APPENDIX 2 Preferred Most Parsimonious Cladogram Found by NONA Cladogram found by NONA using the matrix of Huber (2000) with the additional data in appendix 1 (length ϭ 180; CI ϭ 38; RI ϭ 77). Only clades discussed in the text are numbered. Bold lines indicate Australian taxa. See Relationships for detailed discussion. 144 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 260

APPENDIX 3 Preferred Most Parsimonious Cladogram Found by Pee-Wee Cladogram found by Pee-Wee using the matrix of Huber (2000) with the additional data in appendix 1 (length ϭ 184; CI ϭ 37; RI ϭ 76). Only clades discussed in the text are numbered. Bold lines indicate Australian taxa. See Relationships for detailed discussion.