© The Authors, 2017. Journal compilation © Australian Museum, Sydney, 2017 Records of the Australian Museum (2017) Vol. 69, issue number 1, pp. 1–13. ISSN 0067-1975 (print), ISSN 2201-4349 (online) https://doi.org/10.3853/j.2201-4349.69.2017.1672 Notes on the Ant-mimic Genus Anatea Berland (Araneae: Theridiidae) and Two New Species from Tropical Australia Helen M. Smith, Mark S. Harvey, Ingi Agnarsson and Gregory J. Anderson 1 2 3 4 * 1 Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney NSW 2010, Australia 2 Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC WA 6986; School of Animal Biology, University of Western Australia WA 6009; School of Natural Sciences, Edith Cowan University, Joondalup WA 6009, Australia 3 University of Vermont, Department of Biology, 109 Carrigan Drive, Burlington, VT, 05405-0086; Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC, United States of America 4 Iron Metabolism Laboratory, QIMR Berghofer Medical Research Institute, 300 Herston Road, Herston QLD 4006, Australia Abstract. The taxonomic history of the New Caledonian myrmecomorph spider, Anatea formicaria Berland (Hadrotarsinae: Theridiidae) is summarized, new records are presented and the female is figured for the first time. Two new species provisionally assigned to the genus are described from north-eastern Australia, A. monteithi Smith sp. nov. and A. elongata Smith sp. nov. Some undescribed Anatea species occurring on New Caledonia are shown, and aspects of hadrotarsine anatomy and ant specialization are discussed. Keywords. Anatea formicaria; myrmecomorphy; myrmecophagy; New Caledonia; Queensland; taxonomy Smith, Helen M., Mark S. Harvey, Ingi Agnarsson, and Gregory J. Anderson. 2017. Notes on the ant-mimic genus Anatea Berland (Araneae: Theridiidae) and two new species from tropical Australia. Records of the Australian Museum 69(1): 1–13. https://doi.org/10.3853/j.2201-4349.69.2017.1672 The New Caledonian hadrotarsine spider, Anatea formicaria the “extra” body section (a “petiole”) as well as sometimes Berland, 1927, is thought to be a morphological ant mimic, a postpetiole from constrictions in the cephalothorax or or myrmecomorph (Berland, 1927; Reiskind & Levi, 1967; abdomen and / or colour (Reiskind, 1972), A. formicaria is Cushing, 1997). Myrmecomorphs are especially prevalent in unusual in that the primary modification is to the pedicel, families such as Salticidae and Corinnidae (Cushing, 1997) which is not only elongated, but has an ant-like “node” but there are few in Theridiidae, with only five species in four (Figs 1, 5). Most hadrotarsines have not been thought of as genera listed by Cushing (1997; the sixth species listed appears particularly ant-like, although there are certain aspects of the to be a lapsus). Whilst most myrmecomorph spiders gain anatomy of many species that could aid them hiding among * Corresponding author: [email protected] 2 Records of the Australian Museum (2017) Vol. 69 ant corpses—as recorded for a “Dipoena sp.” regarded as web-building non-mimic by McIver and Stonedahl (1993), a myrmecophile (Cushing, 1997). The adaptations of A. and Cushing (1997) overlooked Berland’s description and formicaria are therefore particularly striking. Associated listed the female as unknown. But as described by Berland behavioural traits that enhance an ant-like appearance, such (1927), and shown here (Figs 5–7), it is similar to the male as running on six legs and holding the front legs aloft like but with reduced scuta. antennae, are common among better known myrmecomorph An ongoing review of Australian Hadrotarsinae has taxa, but we have found no field observations of the behaviour found two new species in northern Queensland that possess of A. formicaria in the literature. certain characters of Anatea, and in particular, they have an Many spiders in the subfamily Hadrotarsinae have been elongate pedicel that is currently diagnostic for the genus. noted to feed on ants. Cushing (2012) lists 10 hadrotarsine The new species we describe are, however, somewhat taxa as myrmecophages (plus “other Euryopis spp”). Indeed, tentatively placed here until further analyses are completed Liu et al. (2016) based on a phylogenetic analysis, found and, hopefully, the unknown male of one species is found. that ant feeding is ancestral for the subfamily, though based Here we figure the correct male of the New Caledonian on relatively sparse taxon sampling. They also found that: Anatea formicaria, and also figure a female of that species (a) subfamily Hadrotarsinae originates about the same time for the first time. We describe the two new species from that ants become noticeably abundant in the fossil record (c. north-eastern Australia as Anatea monteithi sp. nov. and 55–45 Mya); (b) there is a strong correspondence between Anatea elongata sp. nov. We also provide a brief introduction the abundance of ants and the diversity and abundance of to some of the New Caledonian Anatea diversity awaiting hadrotarsines through evolutionary time; and (c) there is a researchers. similar correspondence in modern biodiversity inventories (Liu et al., 2016). Finally, the small, flexible chelicerae with sickle-shaped fangs, suggested by Reiskind & Levi (1967) to Methods possibly indicate an ant-specialized diet, are found in most Specimen examinations, measurements and drawings hadrotarsine taxa. Nevertheless, for the majority of species, were made using an Olympus SZ16 microscope, fitted like A. formicaria, we have no field observations. Our own with graticule and camera lucida. All measurements are observations of Australian hadrotarsines hunting, and with in millimetres. Multiple photographs in different focal prey support the suggestion that many, if not all taxa are planes were taken using a Leica MZ16 microscope fitted primarily myrmecophagous. Since species of Anatea possess with Spot Flex 15.2 Mp camera and layers combined using similar cheliceral modifications to the known hadrotarsine HeliconFocus 5.3 software. Plates were composed using myrmecophages, they may also feed on ants, or similar prey. Adobe Photoshop Elements. Maps were created using Berland came across the specimens he described as ArcGISt software by Esri (www.esri.com). Square brackets Anatea among ants along with some other ant-mimicking enclose notes added to original specimen record information. spider taxa in the collection of myrmecologist Ernest André Specimen preparations for scanning electron microscopy (1838–1911). He surmised that although no doubt André were passed through an alcohol series (75–100 %), critical knew these specimens were spiders, the original collector had point dried and mounted on wire or pins before coating with probably mistaken them for ants whilst collecting. Then, as gold and viewing using a Zeiss Evo LS15 SEM incorporating now, arachnologists recognized that ant mimicking spiders a Robinson backscatter detector. Details of female genitalia occur in several families (and are especially prevalent in were usually drawn from an excised epigynum, briefly Salticidae; Cushing, 1997). However, spiders of several cleared in lactic acid if necessary. genera that we now recognize as being unrelated were then commonly considered to belong together in the Clubionidae. Abbreviations Berland accordingly placed his monotypic genus Anatea there, in subfamily Micariinae, and suggested a relationship All examined specimens are from Australian collections with Micaria Westring, 1851, a genus now included in the as indicated by the following: AM, Australian Museum, Gnaphosidae (World Spider Catalog, 2017). Sydney; ANIC, Australian National Insect Collection, Anatea remained in the Clubionidae until its affiliations Canberra; QM, Queensland Museum, Brisbane; WAM, with Euryopis Menge and Dipoena Thorell were recognized Western Australian Museum, Perth. and it was transferred to Theridiidae (Reiskind & Levi, Morphological abbreviations: AME anterior median 1967). In this paper Reiskind & Levi described a male, which eye; ALE anterior lateral eye; Co conductor; Cy cymbium; they attributed to A. formicaria, although we now realise E embolus; ES embolus spire; FD fertilisation duct; ID this identification was incorrect (see below). However, no insemination duct; MA median apophysis; PME posterior female Anatea has ever been figured despite being included median eye; PLE posterior lateral eye; Sp spermathecae; in the original description by Berland (1927). The female ST subtegulum; T tegulum; TR retrolateral tegulum rim; and of A. formicaria was erroneously characterized as being a TTA theridiid tegular apophysis. Smith et al.: The ant-mimic spider genus Anatea 3 Figures 1–7. Anatea formicaria Berland, (1–4) male, habitus and eyes (AM KS.64090): (1) dorsal; (2) lateral; (3) ventral; (4) prosoma, frontal. (5–7) Female, habitus and opisthosoma (QM S34107): (5) dorsal; (6) lateral; (7) ventral. Scale bars (1, 5) 0.5 mm. 4 Records of the Australian Museum (2017) Vol. 69 Figures 8–11. Anatea formicaria Berland, genitalia, (8, 9) male palpus (AM KS.64090): (8) prolateral; (9) retrolateral. (10. 11) Female (QM S107230): (10) epigynum, ventral (cleared); (11) internal genitalia, dorsal. Scale bars 0.1 mm. Taxonomy Family Theridiidae Sundevall, 1833 Subfamily Hadrotarsinae Thorell, 1881 Anatea Berland, 1927 Anatea Berland, 1927: 53–55. Type species Anatea formicaria Berland by monotypy. Diagnosis. Spiders of genus Anatea exhibit several typical hadrotarsine characters, e.g., sperm duct entering embolus counter-clockwise, cheliceral promargin without teeth, shortened cheliceral base, long slender cheliceral fang, palmate female palpal claw, grouped flat-tipped sensory setae on tarsus I (Agnarsson, 2004). They differ from many other hadrotarsine genera by having only one main pair of spermathecae and carapace not elevated, and differ from all known hadrotarsine genera by the extended and sometimes modified pedicel. Smith et al.: The ant-mimic spider genus Anatea 5 Anatea formicaria Berland (which we have re-identified as a closely related species, “J5”, see Figs 41, 42), they write: “Anatea formicaria is an accurate Figs 1–12, 42 mimic of the small myrmicine ant Chelaner croceiventre (Emery), 2.6 mm long, which has been collected at the same Anatea formicaria Berland, 1927: 55, figs 1–7. locality as the spider. The color pattern (dark brown anterior Not species figured by Reiskind & Levi, 1967: 21, figs 1–6 and light, yellow-brown posterior) of both is quite rare in ants (misidentification, we refer this to sp. “J5”, Fig. 41). and is also found in specimens of Xiphomyrma tenuierius Emery, 2.9 mm. long, and a species of Lordomyrma, 4.8 Diagnosis. Male: From other New Caledonian species mm long, two myrmicine ants found in the same area of rain shown in Figs 38–41, 43 by pedicel and carapace forest (E. O. Wilson, pers. comm.).” The colour description morphology and careful comparison of palp, particularly given above, does not match the specimens we have seen of embolus length. From Australian species, leg I longer than either A. formicaria or “J5”, in which the cephalothorax is leg III, pedicel with “node”, cymbium with extended apex, only slightly darker than the yellow-amber ground colour of ventral abdomen with ventral plates (but note male of A. the abdominal scute (see Figs 41, 42). This difference may elongata sp. nov. is unknown). Female: from undescribed be due to time since preservation or could represent colour New Caledonian species by general morphology (as male) polymorphism within the species “J5”, as is recorded in some and details of genitalia; from Australian species by leg I longer other myrmecomorphs (see examples in Cushing, 1997). than leg III, pedicel with “node”, dorsal scutum present. Berland’s original description of A. formicaria (1927) gives Type material (not examined). 1♂ type, 1♀ cotype, New colouration of cephalothorax, pedicel and abdominal scuta as Caledonia. Found by Berland (dried) in the collection of “light reddish fawn” (“fauve rougeâtre clair”) but this is of ants of Ernest André. Berland does not state what he did minimal help as these were dried specimens of uncertain age. with the specimens. Three of the Anatea samples we have examined each contain a single ant, which we deduce may have been included Other material examined. NEW CALEDONIA: 1♀ QM by collector or sorter of the specimens as a putative model. S34107, Rivière Bleu, Parc.6, 22°05'S 166°40'E, 160 m, 9–22. All three ants are of similar appearance (but may not be iv.1987, L. Bonnet de Larbogne, J. Chazeau & A. & S. Tillier, conspecific) and are from the genus Pheidole. The Pheidole Malaise trap, rainforest sassafras; 1♂ QM S34113, Rivière specimens are of pale golden colour, similar to A. formicaria Bleu, Parc.7, 22°05'S 166°40'E, 170 m, 25.xi–8.xii.1986, L. or our species “J5” (and possibly similar to juveniles of species Bonnet de Larbogne, J. Chazeau & A. & S. Tillier, Malaise “J1”). All these three Anatea species occur together: QM trap; 1♂ 1♀ QM S20728, Mt Koghis, 22°11'S 166°32'E, 500 S10730 and QM S10732 that now contain A. fomicaria and m, 23–24.v.1987, R. Raven, rainforest, general and night species “J5”, respectively, were separated from QM S88025, collection; 1♀, QM S107230, Pic d’Amoa, N slopes, 20°58'S which now contains species “J1”; QM S88026 contains adults 165°17'E, 500 m, 27.xi.2003–30.1.2004, G. Monteith, with and a juvenile of species “J1” and a subadult A. ?formicaria. Pheidole sp. ant, flight intercept trap, SC11482; 1♂ KS.64090, Of note is the size of the ants, which at c. 1.6 mm total length Mt Panie, 20°38'S 164°46'E, 6.vi.1996, M. Moulds; 1♀ WAM are considerably smaller than adult Anatea of either similarly T86527, Col de Rossettes, 21°27'S, 165°28'E, 11.ii.1993, coloured species, and slightly smaller than the sub adult male Harvey, M.S., Platnick, N. I., Raven, R.J., rainforest litter. Anatea spp. included in two of the samples. Size range. Male (n = 3). Carapace range, 0.75–0.80. Total length 2.05–2.15. Female (n = 3). Carapace range, 0.83–0.92. Total length 2.40–2.55. Berland (1927) gives the total length as 2.2 mm for the species. See Berland (1927) for full male description. Distribution. Recorded from mountainous rainforest sites across the main island of New Caledonia (Fig. 12). Notes. The habitus and genitalia of A. formicaria male and female are figured for comparison with the Australian species. Figure 1 is repeated as Fig. 42 to compare with other New Caledonian species. Whilst Berland (1927) admitted his hypothesis that A. formicaria was a myrmecomorph could not be verified, he pointed out that it was found in a collection of ants and that the original collector had probably mistaken it for an ant; unfortunately, he did not (apparently) record which species or genera of ants the spiders were placed with in the collection. However, he comments (p. 53) that the spiders would match species of either Pheidole or Monomorium, both genera being well-represented in the New Caledonian ant fauna: En étudiant les Fourmis de Nouvelle-Calédonie, on trouve même sans trop de peine que l’Araignée se rapprocherait beaucoup d’un Pheidole ou d’un Monomorium, genres qui y sont très bien représentés. Reiskind & Levi (1967) suggest different candidates as Figure 12. Recorded localities of Anatea formicaria Berland in a model for the species they identify as Anatea formicaria New Caledonia. 6 Records of the Australian Museum (2017) Vol. 69 Anatea monteithi Smith sp. nov. Legs: 4312 (holotype female missing both fourth legs but confirmed in female from other material). Legs with sparse Figures 13–24, 31, 35–37 dorsal macrosetae on patellae and tibiae (a few visible in Figs 13, 14), hadrotarsine flat-topped setae (see Agnarsson, 2004) present on distal tarsus I, few on II. Pedicel sclerotized, open Holotype ♀ QM S25842, Mt Formartine South, 10 km N “S” in lateral view, slightly expanded anterior to apex and Kuranda, [16°43'S 145°37'E, Queensland, Australia] 700 constriction at abdominal articulation (Fig. 18). Abdomen m, 23.xi.1990, G. Monteith, G. Thompson, pyrethrum egg-shaped, widest anterior to mid-point (Figs 17, 19), trees & logs. Paratypes. 1♂, WAM T99420, Daintree NP, constricted by sclerotized band around spinnerets; male with Alexandra Range lookout point, 16°14'15"S 145°26'10"E, dorsal scutum covering central abdomen but leaving bare 6.iv.2009, K. Edward, J. Waldock, sieved litter (WT 11); small surround of cuticle in dorsal view (Fig. 13), sclerotized 1♂, AM KS.126452, Black Mountain Rd, start of track to around pedicel and genital plate (Fig. 17); female without Mt Formartine, 16°45'04"S 145°36'21"E, 425 m, 6.xii.2008, scutum (Fig. 18) and only small sclerotized band around G. Milledge, H. Smith, beat, sweep; [right palp on SEM stub pedicel insertion (Fig. 19). One pair of dorsal apodemes, s/1049; right leg I on s/1050]; both Queensland, Australia. prominent in female, embedded in scutum of male; three Other material examined (Queensland, Australia): 1♀ circular ventral plates/apodemes in both sexes (Figs 17, AM KS.116502, Lake Barrine, E of Yungaburra, 17°15'S 19). Genitalia. Male: Cy blunt with two short, broad based 145°38'E, 31.viii.2011, M. Zabka, B. Patoleta; [abdomen tooth-like modified spines (Fig. 21). Palp with Co, TTA, on SEM stub s/1048, legs I, IV on s/1047; epi cleared]. 1 MA (Fig. 35); TR extends ventrally to wrap over the ES, juv. AM KS.7647, Thornton Peak, N of Daintree, 16°10'S Co arises from apicodorsal T and apicolateral tegular rim 145°22'E, Nov. 1975, M. Gray, leaf litter sample (RF survey (Figs 35, 37). TTA large, arising basally, extending beyond site 40). QM 1 juv. (unregistered), Bellenden Ker Ra, 0.5 km tip of Cy. ES tip resting between TTA and Co. MA base S Cable Tower 7, 17°16'S 145°51'E, 500 m, 25–31.x.1981, and basal E conjoined proventrally (Figs 35, 36). Female: Earthwatch/Qld Museum, rainforest, pyrethrum logs, stones, externally with ridges and grooves in lateral profile (Fig. 23), tree trunks. projecting lip on posterior margin of ovoid fossa. Internally (Fig. 22), ID exit fossa posteriorly and almost immediately Etymology. The species is named for Geoff Monteith, in turn anterolaterally, entering spermathecae Sp at posterior recognition of his enormous contributions to the study of end. Fertilisation ducts FD exit adjacent to ID, with terminal Australian invertebrates through his collecting activities for nodule (possibly degenerate second pair of Sp) before the Queensland Museum. terminal part of FD. Diagnosis. From New Caledonian species by leg III longer Measurements. Male. Carapace range, 0.81 to 0.88 (n = than legs I and II; pedicel without “node”; male without 2). WAM T99420. Total length, 2.35; carapace length, 0.88; extended tip to cymbium; from A. elongata sp. nov. by pedicel width, 0.62; height, 0.37; abdomen length, 1.25; width, 0.91; shorter than carapace; female insemination ducts with short pedicel length, 0.50; clypeus height, 0.22. Eyes: AME, 0.10; conjoined section, entering spermathecae posteriorly (Fig. 22). PME, 0.05; ALE, 0.06; PLE, 0.06; Limbs (femur + (patella- tibia) + metatarsus + tarsus = total): leg I, 0.49 + 0.51 + 0.28 Description + 0.24 = 1.51; leg II, 0.44 + 0.54 + 0.25 + 0.29 = 1.51; leg Colour (in alcohol) (Figs 13–19). Cephalothorax, mouthparts, III, 0.50 + 0.56 + 0.24 + 0.33 = 1.63; leg IV, 0.55 + 0.84 + sternum, pedicel and male scutum dark chestnut brown, glossy, 0.33 + 0.36 = 2.08. except pedicel rugose, female abdomen and parts of male Female. Carapace range, 0.86 (n = 2). Holotype QM S25842. abdomen not covered by scutum dark charcoal grey, slightly Total length, 2.88; carapace length, 0.86; width, 0.69; height, paler ventrally, cuticle glossy between sparse but prominent 0.33; abdomen length, 1.68; width, 1.32; pedicel length, 0.56; setal bases; dorsal and ventral abdominal apodemes dark clypeus height, 0.16. Eyes: AME, 0.10; PME, 0.06; ALE, brown, obvious. Legs: (female) coxae and trochanters white 0.07; PLE, 0.06; Limbs (femur + (patella-tibia) + metatarsus on legs III, IV, suffused with brownish black on legs I, II; + tarsus = total): leg I, 0.49 + 0.60 + 0.31 + 0.26 = 1.66; leg femora pale brown laterally, dark dorsally and ventrally II, 0.48 + 0.58 + 0.26 + 0.30 = 1.61; leg III, 0.50 + 0.61 + (legs I, II) or vis. v. (legs III, IV), femora legs III, IV with 0.26 + 0.34 = 1.71; leg IV, missing (but longest in female distal white band extending onto proximal patella, traces of AM KS.116502). same legs I, II; tibiae-tarsi mid-brown proximally to amber brown distally, darkest coloration ventral on anterior legs to Distribution. Northeast Queensland. Recorded from retrolateral on leg IV; male legs similar but darker, stronger rainforests, mainly at higher altitudes, between Thornton coloration. Peak and Bellenden Ker, and on the Atherton Tableland at Yungaburra (Fig. 31). Carapace (Figs 13–15, 18) longer than wide, widest at coxae II, smooth transition to caput; fovea absent; in profile Notes. Without observations of behaviour, it is unclear gently domed, highest at coxae I (male WAM T99420 rather exactly if, or how, ant mimicry is achieved in the Australian flatter than female holotype, other male similar to female); species, but as in New Caledonian species, the long pedicel eye group 3/5 carapace width; AME largest (Figs 14, 15) and overall appearance are ant-like. Relative leg length, and prominent above slightly concave clypeus, clypeus with both III and IV longer than I or II is unusual and may > 2 × AME diameter in male, slightly lower in female. indicate behavioural modifications. Labium bluntly triangular (Fig. 16), cheliceral bases shorter The tropical rainforests of northeastern Queensland than maxillae, flexible with slender, curved fangs. Female occupied by A. monteithi hold an extremely high diversity palpal claw weakly palmate with three or four large teeth. of ants, with over 66 genera recorded in these areas (Antwiki, Smith et al.: The ant-mimic spider genus Anatea 7 Figures 13–19. Anatea monteithi sp. nov. (13–17) male, habitus and detail (WAM T99420): (13) dorsal; (14) lateral; (15) frontal; (16) prosoma, ventral; (17) opisthosoma, ventral. (18, 19) Holotype female (QM S25842): (18) habitus, lateral; (19) opisthosoma, ventral. Scale bars (13, 18) 0.5 mm. 8 Records of the Australian Museum (2017) Vol. 69 Figures 20–24. Anatea monteithi sp. nov. genitalia, (20, 21) male palpus (AM KS.126452): (20) prolateral; (21) ventral. (22–24) Female: (22) internal, dorsal (AM KS.116502); (23, 24) external views (holotype QM S25842): (23) lateral; (24) ventral. Scale bars 0.1 mm. FD, fertilization duct; ID, insemination duct; Sp, spermatheca. 2017). If A. monteithi is indeed an ant mimic, candidate Diagnosis. From New Caledonian species by leg III longer models are found in several myrmecine genera. Examples than legs I and II; pedicel longer than carapace, without of genera with species in an appropriate size range for adult “node”; from A. monteithi sp. nov. by pedicel longer than spiders include Monomorium, Orectognathus, Pheidole, carapace; insemination ducts with long conjoined section, Pristomyrmex and Vombisidris. crossing ventral spermathecae to entry point towards anterior (Fig. 30). Anatea elongata Smith sp. nov. Description Colour (in alcohol). Probably similar to A. monteithi but Figures 25–31 specimen extremely faded, pedicel appears rather reddish- Holotype ♀ ANIC 42 002257, 12 km SSE Heathlands, brown; coxae brown, trochanters white on legs III and IV; 11°51'S 142°38'E [Queensland, Australia], 25.iv–7.vi.1992, otherwise apparently similar to A. monteithi except extent T. McLeod, FIT#2, F.I.T., ANIC 1248, closed forest. of abdominal colour cannot be established as only 1 patch remains. Etymology. The species is named in reference to the pedicel, the longest compared to carapace length so far observed in Carapace (Figs 25, 26, 29) generally similar to female of genus Anatea. A. monteithi; eye group slightly over half carapace width Smith et al.: The ant-mimic spider genus Anatea 9 Figures 25–30. Anatea elongata sp. nov. holotype female (ANIC 42 002257). (25–27) Habitus: (25) dorsal; (26) lateral; (27) ventral. (28) Opisthosoma, ventral. (29) Prosoma, frontal. (30) Epigynum and internal structure, ventral (uncleared). Scale bar (25) 0.5 mm, (30) 0.1 mm. 10 Records of the Australian Museum (2017) Vol. 69 damaging the specimen. Due to the lack of pigment, the external view is sufficient to see the route of the ducts and this is clearly different from A. monteithi. The habitat around the type locality is generally savannah woodland, or “closed forest” on the label, a contrast to the rainforest habitat of A. monteithi. The condition of the specimen is not good enough to determine gloss (except carapace) or be sure of colour. However, if the species is an ant mimic the extreme extension of the pedicel, should facilitate the identification of the model in ant samples taken from the area. Discussion We here describe two new hadrotarsine species (Theridiidae) in the previously monotypic genus Anatea. Species of this genus are remarkable in showing what appears to be detailed ant mimicry that, unlike in most other ant mimics, is achieved by the modification of the pedicel that connects the cephalothorax to the abdomen. Similarities of Anatea fang morphology to Hadrotarsinae that are known to prey on ants may indicate another example of myrmecophagy in the subfamily, and lends support to a potential coevolutionary history of hadrotarsine and ant lineages (Liu et al., 2016). The New Caledonian specimens of A. formicaria figured here have been extracted from mixed Anatea material Figure 31. Recorded localities of Anatea monteithi sp. nov. () and deposited in the Queensland Museum (see Appendix for Anatea elongata sp. nov. () in North East Queensland. details). Many of these samples were collected by Geoff Monteith, as was at least one of the Australian specimens. These few samples alone suggest that many more Anatea species, or relatives, await discovery in New Caledonia, (Fig. 29); eyes and mouthparts similar to A. monteithi. including species without elongate pedicels and sometimes Female palpal claw strongly palmate with at least 8 teeth. with other pedicular modifications that could mimic ants Legs: 43(12). Pedicel sclerotized, almost straight ventrally in subfamilies other than Myrmicinae (Figs 38–41, 43). It in anterior half then arching posteriorly, narrowest anteriorly has also become apparent that the male of Reiskind & Levi and thickest, slightly ridged at mid-point, much longer than (1967) was not in fact A. formicaria, but a rather similar carapace (Figs 25–27). Abdomen as in A. monteithi. Legs species with a slightly shorter embolus and longer, more missing any visible setae but setal bases suggest macrosetae stepped pedicel—both species were found among these probably present as in A. monteithi. samples (Fig. 41). New Caledonia is not the focus of the present Australian study, so we merely flag the potential Genitalia. Externally with fine edge curving anterior to of this fascinating genus for further work there. However, ovoid fossa, but otherwise apparently smooth in profile. it is noteworthy how abundant and diverse Anatea would Internal genitalia not dissected but visible through cuticle appear to be in New Caledonia in comparison to Australia. (Fig. 30), ID exit fossa to posterior, running together for 2/3 One mixed New Caledonia sample in the QM collection of distance to genital groove, then turning away from each was found to contain three different species; another other and simultaneously turning dorsally, crossing ventral included 8 males of one species. Our Australian material, Sp and entering anteroventrally. Sp lying to either side of by contrast, is all singletons and it has proven difficult to fossa in ventral view. FD not visible. find more. As well as the unique female of A. elongata, A. monteithi is represented by only four adults and two Measurements. Female.Total length, 3.04; carapace length, juveniles. Considering the collecting effort put in by Geoff 0.93; width, 0.72; height, 0.38; abdomen length, 1.32; width, Monteith over many years in Queensland compared to a 0.90; pedicel length, 1.06; clypeus height, 0.17. Eyes: AME, few visits to New Caledonia and use of similar collecting 0.09; PME, 0.06; ALE, 0.06; PLE, 0.06; Limbs (femur + techniques, the difference is astounding. Perhaps the species (patella-tibia) + metatarsus + tarsus = total): leg I, 0.52 + we are describing tentatively as Anatea in Australia might 0.60 + 0.30 + 0.26 = 1.68; leg II, 0.50 + 0.60 + 0.28 + 0.30 be utilising a quite different niche and / or behaviour to their = 1.68; leg III, 0.58 + 0.68 + 0.32 + 0.38 = 1.96; leg IV, 0.70 New Caledonian relatives. + 0.98 + 0.42 + 0.42 = 2.52. There are several differences between the New Caledonian Male. Unknown and Australian species that make us question whether Anatea is the correct placement for these new species. First, the Distribution. Northeast Queensland. Recorded only from unusually long rear legs of the Australian species give a leg the type locality on the Cape York Peninsula (Fig. 31). length pattern of 4312. The typical hadrotarsine leg pattern Notes. The single female has lost most of its colour and may is 4123, which is also found in the New Caledonian Anatea not be robust enough to excise the genitalia without severely species. The long leg III is therefore highly unusual and