Zoosyst. Evol. 99 (2) 2023, 307-323 | DOI 10.3897/zse.99.101417 yee BERLIN Kangaraneus, a new genus of orb-weaving spider from Australia (Araneae, Araneidae) Pedro de S. Castanheira!, Volker W. Framenau!** 1 Harry Butler Institute, Murdoch University, 90 South St, Murdoch, Western Australia 6150, Australia 2 Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia, 6986, Australia 3 Department of Invertebrates, Museum of Nature Hamburg - Zoology, Leibnitz Institute for the Analysis of Biodiversity Change (LIB), Martin- Luther-King-Platz 3, 20146 Hamburg, Germany https://zoobank. org/F 1A740F 4-6BC2-44B4-9F 9F-3222C428B23C Corresponding author: Pedro de S. Castanheira (pedro.castanheira@murdoch.edu.au) Academic editor: Danilo Harms Received 3 February 2023 # Accepted 12 April 2023 Published 31 May 2023 Abstract A new Australian genus in the orb-weaving spider family Araneidae Clerck, 1757 is described, Kangaraneus gen. nov., with K. arenaceus (Keyserling, 1886) comb. nov. (New South Wales, Queensland, South Australia, Victoria and Western Australia) as the type species and also including two other species: Kangaraneus amblycyphus (Simon, 1908) comb. nov. (Australian Capital Territory, New South Wales, Queensland, South Australia, Victoria and Western Australia) and K. farhani sp. nov. (Australian Cap- ital Territory, New South Wales, South Australia, Victoria and Tasmania). The new genus is included in the informally termed Australasian ‘backobourkiine’ clade due to the presence of its putative synapomorphies, specifically a single patellar macroseta on the male pedipalp and its median apophysis forming an arch over the radix. It includes medium-sized orb-weaving spiders (total length 3—12 mm) with distinct humeral humps on the almost triangular abdomen. Therefore, within the backobourktiines, it is somat- ically most similar to Novakiella Court & Forster, 1993 but differs considerably in male genitalic characters, including a C-shaped median apophysis with an acute tip. Genitalia are most similar to those in Quokkaraneus Castanheira & Framenau, 2023 from which the new genus differs by the lack of the white colouration and the shape of the abdomen. Key Words Araneus, backobourkiines, systematics, taxonomy Introduction A recent multi-gene molecular study on world-wide orb-weaving spiders in the family Araneidae Clerck, 1757 placed many Australian species in an informally termed clade, the ‘backobourkiines’ (Scharff et al. 2020). The study did not analyse morphological characters in de- tail but proposed two synapomorphies of the male pedi- palp for this clade: a single macroseta on the pedipalp patella — most ‘traditional’ Araneinae Clerck, 1757 have two (Scharff and Coddington 1997) — and the base of the median apophysis forming an arch over the radix. The study by Scharff et al. (2020) spawned several genus-lev- el revisions for the backobourkiines, a dominant Oriental and Pacific clade, that currently includes eleven genera in Australia (Table 1). Two of these genera, Leviana Framenau & Kuntner, 2022 and Quokkaraneus Castan- heira & Framenau, 2022, were placed outside the backo- bourkiines in Scharff et al.’s (2020) preliminary phyloge- netic hypothesis, but as their male pedipalps display the proposed synapomorphies, they are tentatively included in the backobourkiines pending further molecular and morphological studies (see discussions in Framenau and Kuntner (2022) and Castanheira and Framenau (2022)). Copyright Castanheira, P. de S. & Framenau, V. W. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 308 Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia Table 1. Putative genera of the informally termed backobourkiine clade (sensu Scharff et al. 2020) occurring in Australia. Genus Total no. of species; No. of Remarks (World Spider | species in Catalog 2023) Australia Acroaspis Karsch, 1878 5 5 included in Scharff et al. (2020); paraphyletic with respect to Socca; taxonomy unresolved Backobourkia Framenau, Dupérré, 4 4 revised in Framenau et al. (2010); B. thyridota likely senior synonym of Blackledge & Vink, 2010 either B. heroine (L. Koch, 1871) or B. brounii (Urquhart, 1885) pending an examination of type material (whereabouts currently unknown) Carepalxis L. Koch, 1872 i? 5 Taxonomy unresolved; Nearctic species reviewed in Levi (1992) and Ferreira- Sousa and Motta (2022) Hortophora Framenau & Castanheira, 2021 T3 10 revised in Framenau et al. (2021a) Kangaraneus gen. nov. 3 = this study Lariniophora Framenau, 2011 1 1 monotypic (Framenau 201 1) Leviana Framenau & Kuntner, 2022 5 5 revised in Framenau and Kuntner (2022); not within backobourkiines in Scharff et al. (2020), but has putative synapomorphies of backobourkiines Novakiella Court & Forster, 1993 2 2 revised in Framenau et al. (2021b) Plebs Joseph & Framenau, 2012 22 7 revised in Joseph and Framenau (2012) Quokkaraneus Castanheira & Framenau, 1 1 monotypic (Castanheira and Framenau 2022); not within backobourkiines in 2022 Scharff et al. (2020), but has putative synapomorphies of backobourkiines Salsa Framenau & Castanheira, 2022 5 5 revised in Framenau and Castanheira (2022) Socca Framenau, Castanheira & Vink, 2022 12 12 revised in Framenau et al. (2022) dehaantgroup (sensu Yin et al. 1997) 3 1 Parawixia dehaani (Doleschall, 1859) was part of the backobourkiines in Scharff et al. (2020) and not monophyletic with true Parawixia F.O. Pickard- Cambridge, 1904 from the New World The Australian Araneus arenaceus (Keyserling, 1886), originally described from New South Wales and Queensland, and Araneus amblycyphus Simon, 1908 from Western Australia, are morphologically very similar species and both display the proposed male genitalic syn- apomorphies of the backobourkiines as mentioned above. However, they do not display any of the genus-level syn- apomorphies that characterise the backobourkiine genera treated so far (Table 1). They were not included in Scharff et al.’s (2020) or any other recent molecular analysis in- cluding Australian araneids (e.g., Kallal and Hormiga 2018) that could facilitate their phylogenetic placement. Due to their unique morphology, we consider Araneus arenaceus and A. amblycyphus to represent a new genus within the backobourkiines, pending a comprehensive mo- lecular and systematic study of this clade. The aim of this study is to revise this new genus as a testable hypothesis for future systematic work, specifically to elucidate phy- logenetic relationships to or within the backobourkiines. Methods This study is based on the examination of virtually all orb-weaving spider specimens in the major Australian museum collections and overseas collections where his- torical type material is lodged, totalling almost 12,000 records (vials) up to this date. Descriptions and terminology follow recent publica- tions on Australian backobourkiine orb-weaving spiders (e.g., Castanheira and Framenau 2022; Framenau et al. 2022; Framenau and Castanheira 2022; Framenau and Kuntner 2022). The generic and species descriptions of Araneus arenaceus and A. amblycyphus are based on recently collected, well preserved specimens in lieu of original type material. Colour patterns were described based on specimens preserved in ca. 75% ethanol. zse.pensoft.net Microscopic photographs were taken with two different stereo-imaging systems. A setup at the Natural History Mu- seum, Copenhagen (Denmark) allowed taking images with a Nikon D300 digital SLR camera attached via a C-mount adapter to a Leica M16A stereomicroscope. Images of dif- ferent focal plains were stacked with Automontage (vers. 5.02) software from Syncroscopy to increase depth of field. A second set-up at the Harry Butler Institute, Murdoch Uni- versity (Australia) supported taking microscopic images in different focal planes with a Leica DMC4500 digital cam- era mounted to a Leica M205C stereomicroscope and com- bined using the Leica Application Suite X, v. 3.6.0.20104. All photos and plates were edited with Photoshop CC 2020. All measurements are given in millimetres. They were taken with an accuracy of one tenth of a millimetre, ex- cept for the eye and labium measurements, taken with an accuracy of one hundredth of a millimetre. Maps were compiled in the software package QGis v. 3.2.6 Buenos Aires (https://qgis.org/en/site/,; accessed 1 September 2022). Geographic coordinates were extract- ed directly from original labels or the registration data as provided by the museums. When no detailed geograph- ic information was available, localities were estimated based on Google Earth v. 9.1.39.3 (https://earth. google. com/web/ accessed 1 September 2022) to the closest min- ute of Latitude and Longitude. Abbreviations Morphology anterior lateral eyes AME anterior median eyes PLE posterior lateral eyes PME posterior median eyes TL Total length Zoosyst. Evol. 99 (2) 2023, 307-323 Collections AM Australian Museum, Sydney (Australia); CVIC La Trobe University, Bendigo (Australia); MNHN~ Muséum National d’ Histoire Naturelle, Paris (France); MV Museums Victoria, Melbourne (Australia); NHMUK Natural History Museum, London (United Kingdom); QM Queensland Museum, Brisbane (Australia); QVMAG Queen Victoria Museum & Art Gallery, Laun- ceston (Australia) SAM South Australian Museum, Adelaide (Australia); WAM Western Australian Museum, Perth (Australia); ZMB Museum fiir Naturkunde, Zentralinstitut der Humboldt-Universitat, Berlin (Germany); ZMH Zoologisches Institut und Zoologisches Museum, Universitat Hamburg, Hamburg (Germany). Taxonomy Order Araneae Clerck, 1757 Family Araneidae Clerck, 1757 Kangaraneus gen. nov. https://zoobank. org/1 AED6CD8-0E08-4945-83FB-9C 1 EB407717B Type species. Epeira arenacea Keyserling, 1886 (desig- nated here). Etymology. The genus-group name is a compound noun composed of ‘Kang-’ in reference to the Kanga- roo, genus Macropus Shaw, 1790, the renowned Aus- tralian marsupial, and ‘-araneus’, a genus-group name for orb-weaving spiders. The gender of the genus-group name Kangaraneus 1s masculine. Diagnosis. Within the backobourkiines as circumscribed by Scharff et al. (2020; see also Table 1), the male genitalia of Kangaraneus gen. nov. are most similar to those of Quokkaraneus, due to a similar C-shaped median apophysis with a drawn-out, acute tip and the conductor having a basal protrusion (e.g., Castanheira and Framenau 2022; fig. 1C, D). However, Kangaraneus gen. nov. males differ distinctly by the presence of prominent humeral humps on the abdomen (lacking in Quokkaraneus) and the lack of white colouration on the carapace and legs (e.g., Castanheira and Framenau 2022; fig. 1A). Females of Kangaraneus gen. nov. differ from those of Quokkaraneus in somatic characters as described for males above, but also in the general shape of the epigyne scape, which is much wider basally in Quokkaraneus (Castanheira and Framenau 2022; fig. 3C), but not in Kangaraneus gen. nov. Due to the prominent humeral humps on the abdomen, Kangaraneus gen. nov. species are somatically similar to species of Novakiella but have a vastly different genital morphology; specifically, the male pedipalp lacks the greatly enlarged conductor 309 lobe of Novakiella (e.g., Framenau et al. 2021b; figs 1C, 2A). Description. Medium-sized orb-weaving spiders, with males (ca. TL 3.2-6.7) on average smaller than females (ca. TL 6.3—10.4). Carapace longer than wide with cephalic region slightly narrower in males than in females, light or dark brown (e.g., Figs 1A, 3A, 5A, 6A). Fovea longitudinal in males and transverse in fe- males (e.g., Figs 1A, 3A). AME largest in males, row of posterior eyes recurved, lateral eyes almost touching; posterior lateral eyes largest in females and separated from PME by less than their diameter; AME slightly pro- truding from the carapace in both sexes (e.g., Figs 1A, 3A). Sternum slightly longer than wide, brown to beige and sometimes with dusky borders (e.g., Figs 1B, 3B). Labium wider than long, with anterior glabrous pale edge (e.g., Figs 6B, 8B). Maxillae with glabrous light- er antero-mesal section (e.g., Figs 6B, 8B). Chelicerae with four promarginal teeth and three retromarginal teeth (reduced to three and two respectively in K. farhani sp. nov.). Leg formula I > II > IV > III; tibiae II in males stronger than tibiae I and with strong prolateral spines (e.g., Figs 1A, 5A). Abdomen sub-triangular, generally wider than long, with two conspicuous humeral humps, but without specialized setae, condyles or other struc- tures; colour dorsally varying from beige to olive-grey and black, sometimes with distinct folium pattern and large pale spot (e.g., Figs 1A, 3A, 5A). Venter brown or grey with irregular black setae, sometimes with two pale spots (e.g., Figs 1B, 3B, 5B, 9B). Male pedipalp patella with a single macroseta (e.g., Figs 1C, 6C—E, 7A—D); paracymbium hook-shaped (e.g., Figs 1D, 6D, 7A-C, 9D); median apophysis C-shaped, basally strongest and tapering to an acute tip, at ca. half to % length with a prominent bulge (‘central protrusion’), base forming an arch over radix (e.g., Figs 1C—E, 7A—D); radix prominent (e.g., Figs 1C, 2A, 10A); terminal apophysis elongate, membranous, bub- ble-shaped or lanceolate, with a basal lobe and a pointed tip (e.g., Figs 1C, 2A—D, 10OA—D):; basal conductor lobe inconspicuous, terminally spatulate (e.g., Figs 1C, 2A, 6C, 10A); conductor prominent, laterally broad, apically with a protuberance and basally with an elongate pro- trusion tapering to a bifid tip (e.g., Figs 1C—E, 7A—D, 9C-E); embolus strong, basally elongate, rectangular, heavily sclerotised, with a curved, uncapped tip (e.g., Figs 1C, 2A—D, 9C). Epigyne wider than long, with very wide lateral borders encircling a wide and con- vex atrium (Figs 3E, 8C, 11C); central division narrow (Figs 3E, 8E, 11F); scape basally broadest, either taper- ing (Figs 3C, 8C) or truncated (Fig. 11D); spermathecae around half or more than half atrium length, ovoid, cop- ulatory ducts curved, somewhat in a lyre-formation and heavily sclerotised (Fig. 4A—-F). Distribution. All Australian mainland states and Tasmania, excluding Northern Territory. (Fig. 12). Included species. Kangaraneus arenaceus (Keyser- ling, 1886) comb. nov. (type-species), K. amblycyphus (Simon, 1908) comb. nov., and K. farhani sp. nov. zse.pensoft.net 310 Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia Kangaraneus arenaceus (Keyserling, 1886), comb. nov. Figs 1A-E, 2A-D, 3A-E, 4A, B, 5A-C, 12 Epeira arenacea Keyserling 1886: 145-148, plate 12, figs 2, 2a, 3, 3a, 3b. Araneus arenaceus (Keyserling, 1886).- Rainbow 1911: 182; Bonnet 1955: 438. Type specimens. Lectotype of Epeira arenacea Keyserling, 1886 (designated here): female, Sydney (33°53'S, 151°13'E, New South Wales, AUSTRALIA) (NHM_ 1890.7.1.4138), examined. Paralectotypes: 5 males, 3 females, same locality as lectotype (ZMH Rack (1961 )-catalogue 224), examined; 1 male, same locality as lectotype (NHM 1890.7.1.4137), examined; 1 female, same locality as lectotype, misidentified (this terminal lie conductor protuberance ALY conductor apophysis |p is K. amblycyphus comb. nov.) (NHM 1890.7.1.4139), examined. Other material examined. AUSTRALIA — New South Wales ¢ 2 females, 3 juveniles, ‘Allambi’ Tele- graph Point, 31°19'S, 149°15'E (AM_ KS.56967); ¢ | male, Pittwater, 33°38'S, 151°18'E, (AM KS.31675). Queensland ¢ | male, Blackdown Tableland, SW Rock- hampton, 23°50'S, 149°03'E (QM); ¢ 1 female, same lo- cality (QM); ¢ 1 female, Braemar State Forest, 27°13'S, 150°50'E (QM); ¢ 1 male, 2 females, 4 juveniles, Peak Downs, 22°56'S, 148°05'E (ZMH); ¢ 2 females, same lo- cality (ZMH). South Australia « 1 female, Bridgewater, 35°S, 138°46'E (SAM); © 1 female, Cape du Couedic, on road to Flinder Chase Ranger Station, Kangaroo Island, 36°03'S, 136°42'E (SAM); « 1 female, Dudley conductor rotuberance | conductor P \\ median SAA | / apophysis 5 ( / . IN] v fp paracymbium | E . Figure 1. Kangaraneus arenaceus (Keyserling, 1886) comb. nov., male (WAM T64400). A. Dorsal habitus; B. Ventral habi- tus; D. Left pedipalp, ventral view; E. Left pedipalp, retrolateral view; F. Left pedipalp, dorsal view. Scale bars: 2 mm (A, B); 0.2 mm (C—E). Arrow points to the conductor basal protrusion. zse.pensoft.net Zoosyst. Evol. 99 (2) 2023, 307-323 Conservation Park, Kangaroo Island, 35°48'S, 137°51'E (SAM); ¢ 1 female, Kuitpo Forest Reserve, 35°12'S, 138°40'E (SAM); Victoria * 2 females, unknown locality (MV K-10342); ¢ 1 female, Berwick, Quarry Hill, 38°02'S, 145°20'E (QM); ¢ 1 female, 1 juvenile, Spring Gully, 36°48'S, 144°17'E (CVIC 1174). Western Australia e 2 females, south-west Western Australia (no exact lo- cality) (SAM); ¢ | female, Boddington Bauxite Mine, 32°55'S, 116°26'E (WAM 177221); ¢ 1 female, Bus- selton, 33°39'S, 115°20'E (AM KS.131227); ¢ 1 female, Fitzgerald National Park, Twertup, 34°01'S, 119°20'E (WAM 1T75902); *¢ 1 male, Fitzgerald River National Park, on track to Rose Rock, 34°04'S, 119°25'E (WAM T70101); © 1 male, Huntly Mine, 5 km E Banksiadale Dam, 32°39'S, 116°05'E (WAM T64598); ¢ 1 male, same locality (WAM T64400); ¢ 1 male, Jarrahdale (Alcoa) terminal apophysis embolus conductor terminal apophysis basal lobe j conductor lobe fy LY v, basal haematodocha terminal apophysis conductor protuberance eerie nas conductor median ‘ ‘ if SI" [A / /} 7 terminal __ apophysis — median apophysis median apophysis 311 Mine area, 32°20'S, 116°03'E (WAM 148219); ¢ 1 fe- male, Pemberton, 34°26'S, 116°02'E (AM KS.32901),; ¢ 1 male, 1 female, 1 juvenile, Shannon National Park, 34°40'S, 116°22'E (WAM T70165). Diagnosis. Males of Kangaraneus arenaceus comb. nov. are most similar to those of K. amblycyphus comb. nov. as both have pedipalp with a similar median apoph- ysis with a blunt central protrusion and a conductor with an elongated basal protrusion (Figs 1C—E, 2A—D, 6C-E, 7A-D). However, K. arenaceus comb. nov. is identified by a lanceolate terminal apophysis with a more prominent and sclerotised basal lobe, and lacking a distal spine-like tip (present in K. amblycyphus comb. nov.). Kangaraneus arenaceus comb. nov. is the only species in the genus in which females have baso-lateral flaps on the epigyne base (Figs 3E, 4A, B), and in addition can be differentiated embolus conductor median B embolus terminal conductor apophysis Figure 2. Kangaraneus arenaceus (Keyserling, 1886) comb. nov., male expanded left pedipalp (WAM T64598). A. Ventral view; B. Retrolateral view; C. Dorsal view; D. Apical view. Scale bars: 0.2 mm. zse.pensoft.net ah2 Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia central division — scape f atrium baso-lateral flaps E Figure 3. Kangaraneus arenaceus (Keyserling, 1886) comb. nov., female, (WAM T77221). A. Dorsal habitus; B. Ventral habitus; C. Epigyne, ventral view; D. Epigyne, lateral view; E. Epigyne, posterior view. Scale bars: 2 mm (A, B); 0.2 mm (C—E). from K. amblycyphus comb. nov. by the epigyne scape completely covering the atrium (Figs 3C vs 8C). Redescription. Male (based on WAM T64400). Total length 6.7. Carapace 3.0 long, 2.4 wide, light brown, cen- trally darker around fovea (Fig. 1A). Eye diameter AME 0.11, ALE 0.11, PME 0.12, PLE 0.07; row of eyes: AME 0.51, PME 0.42, PLE 1.12. Chelicerae light brown with four promarginal teeth (third largest) and three retromar- ginal teeth of similar size. Legs light brown mottled dark brown, base of femora lighter (Fig. 1A, B). Leg length of segments (femur + patella + tibia + metatarsus + tarsus = total length): I—- 3.94 16+3.2+2.8 + 1.2 = 12.7, Il —3.14+154+244+23+4+1.0=10.3, Il-11+09+1.4 +1.0+06=5.0,IV—18+0.9+15+1.8+0.8=6.8. Labium 0.47 long, 0.35 wide, light brown; maxillae light zse.pensoft.net brown, both beige anteriorly (Fig. 1B). Sternum 1.3 long, 1.0 wide, light brown with smooth darker radial shading (Fig. 1B). Abdomen 3.5 long, 3.6 wide, triangular with two conspicuous and pointed shoulder humps; dorsum with pale beige background, large olive-grey patch in cardiac area and irregular folium with olive-grey spots, laterally pale beige with dark streaks (Fig. 1A); venter same background colour as dorsum and with sparse gua- nine spots (Fig. 1B). Pedipalp (Figs 1C—E, 2A—D) length of segments (femur + patella + tibia + cymbium = to- tal length): 0.6 + 0.4 + 0.2 + 1.0 = 2.2; paracymbium strong, hook-like and curved apically; median apophy- sis C-shaped, terminating in an acute tip, central protru- sion small; radix strong; terminal apophysis elongated, membranous and lanceolate, bearing a large, sclerotised, Zoosyst. Evol. 99 (2) 2023, 307-323 copulatory duct copulatory duct “a spermathecae copulatory duct x copulatory % duct ee spermathecae copulatory duct SS copulatory yw duct > E “a fertilizatio Mae | J : a ducts 313 * . em spermathecae spermathecae a | ” ; ea | “spermathecae Figure 4. Kangaraneus gen. nov. female epigynes, cleared. A. K. arenaceus (Keyserling, 1886) comb. nov., ventral view (WAM 177221); B. K. arenaceus (Keyserling, 1886) comb. nov., dorsal view (WAM 177221); C. K. amblycyphus (Simon, 1908) comb. nov., ventral view (AM KS.131276); D. K. amblycyphus (Simon, 1908) comb. nov., dorsal view (AM KS. 131276); E. K. farhani Sp. nov. ventral view (AM KS.131278); F. K. farhani sp. nov., dorsal view (AM KS. 131278). Scale bars: 0.2 mm. round projection, ending in an acute tip; conductor large, round and laterally projected, bearing a hook-like apical protuberance and an elongated basal protrusion tapering to a bifid tip; basal conductor lobe inconspicuous; em- bolus prominent, elongate, heavily sclerotised with a curved tip. Female (based on WAM 177221): Total length 5.6. Carapace 2.5 long, 2.4 wide; beige, cephalic area light brown (Fig. 3A). Eye diameter AME 0.11, ALE 0.10, PME 0.09, PLE 0.09; row of eyes: AME 0.46, PME 0.38, PLE 1.23. Chelicerae beige, with four promarginal teeth (apical and third largest) and three retromarginal teeth of similar size. Legs beige, patellae and tibiae apically slight- ly darker (Fig. 3A, B). Pedipalp length of segments (femur + patella + tibia + tarsus = total length): 0.6 + 0.4 + 0.4 + 1.2=2.6. Leg length of segments (femur + patella + tibia + metatarsus + tarsus = total length): 1—-2.9+14+24+2.3 +1.1=10.1,0-2.441.142.1421+09=8.6, 11-1.4+ 0.9+1.04+1.0+0.8=5.1, [V—1.8+1.0+15+1.7+1.0 = 7.0. Labium 0.25 long, 0.60 wide, beige; maxillae beige (Fig. 3B). Sternum 1.5 long, 1.2 wide, beige (Fig. 3B). Abdomen 3.5 long, 3.2 wide; dorsum of same colour back- ground as male but almost completely covered greyish, laterally grey (Fig. 3A); venter grey with a few guanine zse.pensoft.net 314 Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia Figure 5. Kangaraneus arenaceus (Keyserling, 1886) comb. nov., variations. A. Male dorsal habitus (WAM T64598); B. Paralecto- type male ventral abdomen (NHM 1890.7.1.4137); C. Lectotype female ventral abdomen (NHM 1890.7.1.4138). spots (Fig. 3B). Epigyne base oval, wider than long, with wide lateral borders, atrium convex, barely visible in ven- tral view (Fig. 3C); central division narrow, baso-lateral flaps present (Figs 3E, 4A); scape elongate-triangular ex- tending over the posterior epigyne border, basally broadly rounded and slightly wrinkled, almost straight in lateral view, tip flattened without terminal pocket (Fig. 3C, D); spermathecae elongate-ovoid, copulatory ducts sinuous and terminating at base of scape (Fig. 4A, B). Variation. Males total length 4.5—6.7 (n = 6); females total length 5.6—8.7 (n = 6). The colouration of K. arena- ceus comb. nov. males can vary considerably, with some specimens having a dark brown carapace and a black abdo- men with a large white patch tn the cardiac area (Fig. 5A). Examined females are less variable in colouration. Both males and females may have a pair of white spots ventral- ly on the abdomen, a character mentioned in the original description (see Keyserling 1886) (Fig. 5B, C). Remarks. The syntypes of Epeira arenacea Key- serling, 1886 are deposited at the ZMH and NHM. A female paralectotype (NHM 1890.7.1.4139) was mis- identified by Keysersing (1886); this is K. amblycyphus comb. nov. To unequivocally fix the species-group name of K. arenaceus comb. nov., we here designate a female collected in Sydney (NHM 1890.7.1.4138) as the lecto- type of the species. We do not consider the specimens from Peak Downs (Queensland) at the ZMH syntypes of K. arenaceus comb. nov. as Keyserling’s type locality is “Sydney. Museum Godeffroy”. Life history and habitat preferences. Specimens of K. arenaceus comb. nov. have mainly been found from zse.pensoft.net October to January, with only a few specimens collected in the colder months like June and August. Therefore, the species seems to be largely spring to early summer mature. Habitat descriptions on the collection labels include “cypress and brigalow” and “in low bush”. Two labels indicated that the species falls prey to wasps (“in wasp nest” and “mud dauber wasp nest’). Distribution. Kangaraneus arenaceus comb. nov. has been found in eastern New South Wales and Queensland, the Fleurieu Peninsula and Kangaroo Island in South Australia, Victoria and south-western Western Australia (Fig. 12). Kangaraneus amblycyphus (Simon, 1908), comb. nov. Figs 6A-E, 7A-D, 8A-E, 12 Araneus amblycyphus Simon 1908: 427. Araneus amblycyphus Simon, 1908.- Rainbow 1911: 182; Bonnet 1955: 432. Type specimens. Lectotype of Araneus amblycyphus Simon, 1908 (designated here): 1 female Busselton (33°38'S, 115°20'E, Western Australia, AUSTRALIA) (MNHN 13309), examined. Paralectotype: 1 juvenile, same locality (ZMB 13485), examined. Other material examined. AUSTRALIA— Australian Capital Territory * 1 female, Corin Dam, 57 km W of Canberra, 35°34'S, 148°50'E (AM KS.32942): New South Wales ¢ | female, Brooklana, W of Dorrigo, 30°16'S, 152°S1'E (AM KS.34050); * 1 male, Hornsby, Waitara Zoosyst. Evol. 99 (2) 2023, 307-323 < s conductor protuberance terminal embolus, conductor conductor / median apophysis © iJ D conductor SS conductor protuberance conductor median apophysis ~ paracymbium paracymbium le Figure 6. Kangaraneus amblycyphus (Simon, 1908) comb. nov., male (AM KS.131277). A. Dorsal habitus; B. Ventral habitus; C. Left pedipalp, ventral view; D. Left pedipalp, retrolateral view; E. Left pedipalp, dorsal view. Scale bars: 2 mm (A, B); 0.2 mm (C—E). Arrow points to the conductor basal protrusion. Creek, 33°42'S, 151°0S'E (AM KS.91125); ¢ 1 female, same data (AM KS.91105); ¢ 1 female, Howes Valley, 32°51'S, 1S50°S1'E (AM KS.34043); ¢ 1 male, Jamberoo Mountain, 34°40'S, 150°43'E(AM KS.54098); «2 females, Scheyville, 33°36'S, 150°53'E(AM KS.98752); « 1 female, same data (AM KS.131276); « 1 female, Sydney, 33°53'S, 151°13'E (NHM 1890.7.1.4139; paralectotype of K. are- naceus comb. nov.); ¢ 1 female, 3 juveniles, Wilson River Flora Reserve, Bellangry State Forest, 31°18'S, 152°29'E (AM KS.9666); Queensland « | female, Braemar State Forest, 27°13'S, 150°50'E (QM); * 1 male, same locali- ty (QM); ¢ 1 male, 6 juveniles, same locality (QM); 1 female, 1 juvenile, Camira, 27°38'S, 152°55'E (QM S2462); * 5 females, same locality (QM); * 2 males, Malanda, SW, Lot 2, Meragallan Road, 17°25'S, 145°32'E (AM KS.86086); * 1 male, same data (AM KS.131277); e 1 male, Mt Lewis, 16°35'S, 145°17'E (QM); © 1 male, Mulgowie, 27°43'S, 152°22'E (QM); ¢ 1 female, Tweed River, 28°18'S, 153°27'E (AM KS.32649); South Aus- tralia * 2 females, Dudley Conservation Park, Kangaroo Island, 35°48'S, 137°51'E, (SAM); Victoria « 1 female, German Creek, near Bright, 36°43'S, 147°02'E (QM); ¢ 2 females, Snobs Creek, 37°15'S, 145°52'E (MV K-9440). Western Australia: * 2 males, 5 females, Stirling Range National Park, Moingup Spring, 34°24'S, 118°06'E (WAM 174424). Diagnosis. Male K. amblycyphus comb. nov. can be distinguished from those of the very similar K. arenaceus zse.pensoft.net 316 Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia embolus — terminal apophysis conductor _ ———_ _ basal haematodocha ‘paracymbium terminal —— apophysis embolus conductor Conductor protuberance LAN conductor protuberance en median apophysis | 8 — terminal apophysis embolus — median apophysis terminal embolus — apophysis conductor median apophysis D Figure 7. Kangaraneus amblycyphus (Simon, 1908) comb. nov., male expanded left pedipalp (AM KS.131277). A. Ventral view; B. Retrolateral view; C. Dorsal view; D. Apical view. Scale bars: 0.2 mm. comb. nov. by the double central protrusion on the medi- an apophysis, the bubble-shaped terminal apophysis, and the shorter and thicker basal protrusion of the conductor with a lamellar apical protuberance (Figs 6A—E, 7A—D). Females of K. amblycyphus comb. nov. can be differenti- ated from those of other Kangaraneus gen. nov. species by the epigyne scape not entirely covering the atrium (Fig. 8A, C). Redescription. Male (based on AM KS.131277). Total length 5.8. Carapace 2.6 long, 2.4 wide, light brown, anteriorly darker (Fig. 6A). Eye diameter AME 0.12, ALE 0.11, PME 0.15, PLE 0.13; row of eyes: AME 0.54, PME 0.46, PLE 1.0. Chelicerae light brown, basally mottled dark brown; with four promarginal teeth (second from base largest) and three retromarginal teeth of similar zse.pensoft.net size. Legs light brown, mottled in dark brown mainly on femora (Fig. 6A, B). Leg length of segments (femur + patella + tibia + metatarsus + tarsus = total length): [—2.9 a ASR OS 2 eh de 2 ahi a Nee OA ee 1.1=8.3, WI-1.0+0.9+1.2+1.0+0.8=4.9, IV—1.6+ 1.1+15+1.4+0.8=6.4. Labium 0.30 long, 0.43 wide, yellowish brown; maxillae yellowish brown (Fig. 6B). Sternum 1.0 long, 0.9 wide, light brown with black radial shading (Fig. 6B). Abdomen 2.8 long, 2.8 wide, oval with inconspicuous shoulder humps; dorsum with beige back- ground and large olive-grey, irregular folium bordered anteriorly and posteriorly by brown bands, laterally beige with dark streaks (Fig. 6A); venter olive-grey, laterally with two pale bands (Fig. 6B). Pedipalp (Figs 6C—E, 7A— D) length of segments (femur + patella + tibia + cymbium Zoosyst. Evol. 99 (2) 2023, 307-323 atrium C D oy ie ‘ central division E Figure 8. Kangaraneus amblycyphus (Simon, 1908) comb. nov., female, (AM KS.131276). A. Dorsal habitus; B. Ventral habitus; C. Epigyne, ventral view; D. Epigyne, lateral view; E. Epigyne, posterior view. Scale bars: 2 mm (A, B); 0.2 mm (C—E). = total length): 0.7+ 0.2 + 0.3 + 1.1 =2.3; paracymbium short, hook-like and slightly curved apically; median apophysis C-shaped, terminating in an acute tip, central protrusion accompanied by smaller bulge apically; radix comparatively slim; terminal apophysis elongate, slightly inflated and bubble-shaped, terminating in an acute spine- like tip; basal conductor lobe large; conductor prominent, bearing a strong protrusion tapering to a bifid tip; em- bolus short and elongate, heavily sclerotised with its tip curved apically. Female (based on AM KS.131276): Total length 5.8. Carapace 2.7 long, 2.6 wide; dark brown, (Fig. 8A). Eye diameter AME 0.09, ALE 0.08, PME 0.10, PLE 0.07; row of eyes: AME 0.46, PME 0.41, PLE 1.15. Chelicerae dark brown, four promarginal teeth (second basal largest) and three retromarginal teeth of similar size. Legs brown with yellowish setae (Fig. 8A, B). Pedipalp length of segments (femur + patella + tibia + tarsus = total length): 0.9 + 0.5 + 0.7+ 0.8 =2.9. Leg length of segments (femur + patella + tibia + metatarsus + tarsus = total length): [-— 3.4 + 1.5 +2.84+22+1.0=10.9,11-3.44+144+2.74+22+11= 10.8, WI-1.7+0.9+1.44+1.0+0.7=5.7,1V-2.4+1.1+ 1.54+1.4+0.8=7.2. Labium 0.22 long, 0.49 wide, beige; maxillae beige (Fig. 8B). Sternum 1.3 long, 1.2 wide, dark brown, with yellow streaks (Fig. 8B). Abdomen 3.9 long, 5.4 wide; dorsum with two conspicuous shoulder humps, uniformly beige with indistinct darker line between hu- meral humps (Fig. 8A); venter dark olive grey (Fig. 8B). Epigyne base almost circular, slightly wider than long, with large and elevated borders (Fig. 8C); central division zse.pensoft.net 318 Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia narrow (Fig. 8E); scape elongate, longer than the epigyne base, with broad wrinkled base (Fig. 8C, D); spermathe- cae oval, almost as long as half of the epigyne length, cop- ulatory ducts C-shaped and heavily sclerotised, copulato- ry openings near the base of scape (Fig. 4C, D). Variation. Males total length 3.2—-5.8 (n = 4); females total length 3.5-6.0 (n = 7). There was little colour vari- ation in male K. amblycyphus comb. nov. Some female specimens had a large white patch completely cover- ing the ventral part of abdomen and/or a rounded white patch 1n the cardiac area, similar to males of K. arenaceus comb. nov. and K. farhani sp. nov. Remarks. There are only two syntypes for this spe- cies, one female deposited in Paris (MNHN 13309) and a juvenile deposited in Berlin (ZMB 13485), both from the type-locality of Busselton (Western Australia). We here designate the female as the lectotype of this species as the juvenile cannot be unequivocally identified and may be K. arenaceus comb. nov., which also occurs in Western Australia. Life history and habitat preferences. Mature spec- imens of K. amblycyphus comb. nov. were mainly col- lected between September and December with few specimens collected in February and April; the species therefore appears to be spring to early summer mature. Habitat descriptions on the labels include “cypress and brigalow” and “Callitrus”’; the species was also found in a “wasp nest”. Distribution. The distribution of K. amblycyphus comb. nov. is very similar to that of K. arenaceus comb. nov. although it has been found much further north in Queensland. It occurs in all Australian states except Northern Territory and Tasmania (Fig. 12). Kangaraneus farhani sp. nov. https://zoobank.org/CDODBA93-E176-468F-B6C0-238ED30A 6FFO Figs 9A-E, 10A—D, 11A-F, 12 Type-material. Holotype male, Frankston (38°09'S, 145°08'E, Victoria, AUSTRALIA), 25 September 1947 (MV K-9854). Other material examined. AUSTRALIA — Austra- lian Capital Territory * 2 females, Corin Dam, 35 km W Canberra, 35°34'S, 148°50'E(AM KS.33511). New South Wales « 1 female, Barrington Tops, 32°01'S, 151°29'E (MV K-9814); * 2 males, Jamberoo Mountain, 34°40'S, 150°43'E (AM KS.54090, KS.56902); * 2 females, same locality (AM KS.54096, KS.56898); * 1 female, Jeno- lan Caves, 33°49'S, 150°O1'E (AM KS.32648); « 2 fe- males, New England National Park, 30°29'S, 152°30'E (QM); * 1 female, Newnes State Forest, Sunnyside Road, 1.6 km from Blackfellows Hand Road, 33°22'S, 150°11'E (AM KS.93211). South Australia ¢ 1 female, Mylor, 35°03'S, 138°46'E (SAM); ¢ 1 female, Ravine des Casoars, Kangaroo Island, 35°48'S, 136°45'E (SAM). Tasmania ¢ | female, King Island, 39°53'S, 143°54'E (MV _ K-9468); * 3 females, Launceston, 41°27'S, zse.pensoft.net 147°10"E (AM KS.28590); ¢ 1 female, same locality (AM KS.28697); © 1 male, 2 females, same locality (AM KS.28545); ¢ 1 female, same locality (AM KS.131278); e | female, Lees Paddocks, 41°50'S, 146°06'E (QVMAG 13:0529); * 2 females, Lilydale, 41°15'S, 147°13'E (AM KS.8524); © 1 female, Maggs Mountain, 41°44'S, 146°10'E (QVMAG 13:0634); * 1 female, Picton area, 43°09'S, 146°38'E (MV K-10379); © 1 male, Ringarooma River, Gladstone, 40°57'S 148°01'E (QM §90595); ¢ 1 female, Risdon, 42°49'S, 147°19'E (AM KS.28839); e 1 female, 2 juveniles, Southport, 43°25'S, 146°58'E (AM KS.28870); ¢ 1 female, Stanley, 1 Ford St, 40°46'S, 145°17'E (QVMAG 13:0582); ¢ 1 female, same locali- ty (QVMAG 13:0584); * 1 female, Strahan, 42°09'S, 145°19'E (AM KS.34086). Victoria * 1 female, no exact locality (MV K-10377); © 1 female, Camberwell, 37°50'S, 145°4'E (MV K-10375); ¢ 2 females, Frankston, 38°09'S, 145°08'E (MV _ K-10378); © 1 female, Macclesfield, 37°53'S, 145°29'E (MV K-10376); © 3 females, Narracan, 38°15'S, 146°13'E (MV K-10374). Etymology. The specific name is a patronymic in hon- our of Farhan Bokhari, a colleague at the Harry Butler Institute (Murdoch University). Diagnosis. Male K. farhani sp. nov. can be distin- guished from the other two species of the genus by the central protrusion of the median apophysis being much longer, somewhat blade-like and heavily sclerotised (Figs 9C, D, 10A, B). Females can be differentiated from the other two Kangaraneus gen. nov. species by the trun- cated tip of the epigyne scape (tapering in the other two species) (Fig. 11D—-F). Kangaraneus farhani sp. nov. is the only species in which females were found with a broken off scape exposing a distinct, shallowly V-shaped posterior edge of the atrium (Fig. 10C). Description. Male (based on holotype MV K-9854). Total length 5.8. Carapace 3.6 long, 3.1 wide, light brown, slightly lighter in cephalic area (Fig. 9A). Eye diameter AME 0.23, ALE 0.18, PME 0.18, PLE 0.16; row of eyes: AME 0.64, PME 0.43, PLE 1.17. Chelicerae light brown, basally mottled dark; with three promarginal teeth (me- dian largest) and two retromarginal teeth of similar size. Legs light brown, patched in beige on the base of femur (Fig. 9A, B). Leg length of segments (femur + patella + tibia + metatarsus + tarsus = total length): I - 3.9 + 1.3 2.6.29 P= 11 Oh ee 1 323.24 22 410 = 11.4, 101-2.2+08+16+10+05 =6.1,IV—3.2 + 0.7 + 1.8 + 2.0 + 1.0 = 8.7. Labium 0.39 long, 0.52 wide, light brown; maxillae light brown (Fig. 9B). Ster- num 1.6 long, 1.2 wide, light brown with black contour (Fig. 9B). Abdomen 3.1 long, 3.1 wide, with conspicuous humeral humps; dorsum olive-grey, with short, pale lon- gitudinal patch anteriorly and a dark, heart-shaped patch posteriorly, demarcated by a pale line (Fig. 9A); venter olive-brown with irregular light streaks (Fig. 9B). Pedi- palp (Figs 9C—E, 10A—D) length of segments (femur + patella + tibia + cymbium = total length): 0.7+ 0.4 + 0.6 + 1.3 =3.0; paracymbium strong and hook-like, curved api- cally; median apophysis C-shaped, tapering to an acute Zoosyst. Evol. 99 (2) 2023, 307-323 A terminal apophysis terminal apophysis conductor - conductor apophysis C | bD 319 conductor protuberance a ay ™ median © apophysis paracymbium | cE Figure 9. Kangaraneus farhani sp. nov., male holotype (MV K-9854). A. Dorsal habitus; B. Ventral habitus; C. Left pedipalp, ven- tral view; D. Left pedipalp, retrolateral view; E. Left pedipalp, dorsal view. Scale bars: 2 mm (A, B); 0.2 mm (C-—E). tip, central protrusion elongate, heavily sclerotised; radix canoe-shaped; terminal apophysis elongate, lanceolate, tapering to an acute tip; basal conductor lobe conspicu- ous, heavily sclerotised; conductor triangular, bearing a membranous apical protuberance and a short basal pro- trusion with a bifid tip; embolus heavily sclerotised with its tip curved apically. Female (based on AM KS.54096): Total length 10.4. Carapace 3.7 long, 3.6 wide; colouration as in male (Fig. 11A). Eye diameter AME 0.13, ALE 0.11, PME 0.12, PLE 0.10; row of eyes: AME 0.51, PME 0.43, PLE 1.28. Chelicerae brown, four promarginal teeth (apical and third largest) and three retromarginal teeth of similar size. Legs light brown with covered in setae (Fig. 11A, B). Pedipalp length of segments (femur + pa- tella + tibia + tarsus = total length): 1.3+0.5+0.8+ 1.7 = 4.3. Leg length of segments (femur + patella + tibia + metatarsus + tarsus = total length): 1-4.4+2.0+3.8 + 3.0+ 1.7= 14.9, 1-3.9+ 1.8+3.3+2.6+ 1.4 = 13.0, Thee ee. Geral ses Ole I — 338. + Le +2.2+2.3+1.2 = 10.8. Labium 0.56 long, 0.89 wide, light brown; maxillae light brown (Fig. 11B). Sternum 2.0 long, 1.8 wide, colouration as in male (Fig. 11B). Abdomen 7.9 long, 6.7 wide; dorsum with two conspic- uous humeral humps, golden brown with brown band between humeral humps (Fig. 11A); venter olive-grey with indistinct pale lateral bands (Fig. 11B). Epigyne base rounded, wider than long, with large and elevated borders including a V-shaped posterior rim (Fig. 11C, D); atrium convex (Fig. 11C, E, F); central division wide (Fig. 11F); scape slightly longer than wide, lon- ger than the epigyne base, with rounded bulged base, zse.pensoft.net 320 embolus terminal — apophysis conductor terminal me — Apophysis basal lobe \ PM ‘ 4 \ etn median apophysis basal ~ | haematodocha A terminal apophysis a -paracymbium conductor median apophysis terminal apophysis basal lobe Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia terminal apophysis conductor 4 _ basal haematodocha ee SN Wembolus median \ - apophysis ’ -—. B aati - Z = le median apophysis Figure 10. Kangaraneus farhani sp. nov., male expanded right pedipalp (AM KS.56902). A. Ventral view; B. Retrolateral view; C. Dorsal view; D. Apical view. Scale bars: 0.2 mm. apically truncated (Fig. 11D, E); spermathecae oval, located basally on the genital plate, copulatory ducts short (Fig. 4E, F). Variation. Males total length 5.6—5.8 (n = 2); females total length 6.3-10.4 (n= 8). Males display similar colour variations to K. arenaceus comb. nov. with one specimen having a large white patch in the cardiac area. Females do not display much colour variation, but half had the epigyne scape broken off. Life history and habitat preferences. Mature speci- mens of K. farhani sp. nov. were collected between July and early December, except for one female collected in February. The species therefore seems to be largely win- ter- to spring mature. Kangaraneus farhani sp. nov. generally appears to build its web low in grass, with original labels reading zse.pensoft.net “among grass in garden”, “garden”, “orb-web near ground”, “in grass at side of creek”. Distribution. Kangaraneus farhani sp. nov. occurs in south-eastern mainland Australia (Australian Capital Ter- ritory, New South Wales, South Australia, Victoria) and Tasmania (Fig. 12). Discussion We here consider Kangaraneus gen. nov. to be the twelfth genus within the backobourkiines sensu Scharff et al. (2020) due to the presence of a single patellar macroseta and the median apophysis forming an arch of the radix (Table 1). However, this placement remains ambiguous. Genital morphology, in particular that of the male pedi- Zoosyst. Evol. 99 (2) 2023, 307-323 atrium 321 central division F Figure 11. Kangaraneus farhani sp. nov., female. A. Dorsal habitus (AM KS.54096); B. Ventral habitus (AM KS.54096); C. Broken epigyne, ventral view (AM KS.54096); D. Epigyne, with scape, ventral view (AM KS. 131278); E. Epigyne, lateral view (AM KS.131278); F. Epigyne posterior view (AM KS. 131278). Scale bars: 2 mm (A, B); 0.2 mm (C-—F). palp, is most similar to that of Quokkaraneus necopinus (Keyserling, 1887), but that species was not recovered in the backobourkiines in Scharff et al. (2020); 1t was basal in their ARA-clade with the exception of ‘caerostrines’, phylogenetically far removed from the backobourkiines. However, Scharff et al. (2020; p. 16) stressed the prelim- inary aspect of their phylogenetic hypothesis: “Little can be concluded other than araneid phylogeny remains a work in progress to be pursued with more data and more taxa.” The same is true for the backobourkiines and their putative members (Table 1). More comprehensive phy- logenetic analyses are required to elucidate their rela- tionships and interpret the evolution of morphological characters, including genitalia. The distribution patterns of K. arenaceus comb. nov. and K. amblycyphus comb. nov. are very similar, and there are several localities where both species were found together, even simultaneously (Busselton, Western Aus- tralia; Braemar State Forest, Queensland; Dudley Conser- vation Park, South Australia; Sydney, New South Wales). This made initial matching of sexes difficult. However, as multiple males and females of each species as diagnosed here were found together in several localities (Shannon and Sterling Range National Parks, Western Australia; Hornsby, Waitara Creek, New South Wales; Braemar State Forest, Queensland), we consider the results of our taxonomic revision the most likely match for males and females of K. arenaceus comb. nov. and K. amblycyphus zse.pensoft.net 322 Castanheira, P. de S. & Framenau, V. W.: Kangaraneus, a new genus of Araneidae from Australia 115°0'E 120°0'E 125°0'E 130°0'E $0.91 $0.02 $0.92 $.0.0€ S.0.S¢ O Kangaraeus arenaceus comb. nov. 3 Kangaraeus amblycyphus comb. nov. © Kangaraeus farhani sp. nov. S.0.0P 0 150 300 450 600 km 115°0°'E 120°0'E 125°0'E 130°0'E 135°0'E Teaittory is ie South 135°0'E 140°0'E 145°0°'E 150°0°E 155°0°'E 15°0'S 20°0'S Queansiancd . 25°0'S 30°0'S 35°0'S 40°0'S 140°0'E 145°0'E 150°0’E 155°0'E Figure 12. Distribution records of Kangaraneus comb. nov. species. comb. nov. Our species diagnoses have been compiled based on best evidence and by arachnologists highly ex- perienced in araneoid taxonomy. However, there remains the remote possibility that either males and females are matched incorrectly or, alternatively and even less likely, that male and female genitalia in these species are highly variable and both represent the same species. Future mo- lecular analyses could confirm the matching and there- fore the distribution patterns of these species. Acknowledgements We thank (in no particular order) Peter Lillywhite, Joseph Schubert, Catriona McPhee, Ken Walker, Richard March- ant, Simon Hinkley (NMV), Robert Raven (retired), Owen Seeman, Jeremy Wilson, Michael Rix (QM), John Douglas (retired) (QVMAG), David Hirst (retired), Mat- thew Shaw (SAM), Graham Milledge (retired), Helen Smith (AM), and Mark Harvey and Julianne Waldock (WAM), for the loan of specimens in their care and/or assistance when visiting their respective institutions. We also thank Christine Rollard and Yvan Montardi for shar- ing images of the K. amblycyphus comb. nov. lectotype. Janet Beccaloni and Nikolaj Scharff hosted both authors at a number of occasions in London and Copenhagen, respectively. Nikolaj also provided in-depth insights into araneid phylogeny and systematics. zse.pensoft.net Funding for revisions of the Australian Araneidae was provided by the Australian Biological Resources Study (ABRS) (grant no. 205-24 [2005-2008] to VWF and N. Scharff and grant number 4-EHPVRMK [2021-2023] to VWE, PSC, N. Scharff, D. Dimitrov, A. Chopra and R. Baptista). Additional funding was provided by a Synthe- sis project grant (GB-TAF-TA3-008) to PdSC to visit the NHMUK in London. References Bonnet P (1955) Bibliographia araneorum. Analyse méthodique de toute la littérature aranéologique jusqu’en 1939. Tome II. Systéma- tique des araignées (Etude par ordre alphabétique) [1° partie: A—B]. Douladoure, Toulouse, 1-918. Castanheira P de S, Framenau VW (2022) Quokkaraneus, a new mono- typic genus of Australian orb-weaving spider (Araneae, Araneidae). Australian Journal of Taxonomy 10: 1-9. https://doi.org/10.54102/ ajt.a7o0q2 Castanheira P de S, Framenau VW (2023) Abba, a new monotypic ge- nus of orb-weaving spiders (Araneae, Araneidae) from Australia. Evolutionary Systematics 7(1): 73-81. https://doi.org/10.3897/evol- syst.7.98015 Ferreira-Sousa L, Motta PC (2022) Diagnostic notes on the spider orb-weaving genera Carepalxis and Ocrepeira (Araneae: Aranei- dae), with description of three new species from Central Brazil. Zoo- taxa 5087(2): 389-399. https://doi.org/10.11646/zootaxa.5087.2.9 Zoosyst. Evol. 99 (2) 2023, 307-323 Framenau VW (2011) Lariniophora, a new monotypic orb-weaving spi- der genus from Australia (Araneae: Araneidae: Araneinae). Records of the Western Australian Museum 26(2): 191-201. https://doi. org/10.18195/issn.0312-3162.26(2).2011.191-201 Framenau VW, Castanheira P de S (2022) Revision of the new Australasian orb-weaving spider genus Salsa (Araneae, Aranei- dae). ZooKeys 1102: 107-148. https://doi.org/10.3897/zook- eys.1102.82388 Framenau VW, Kuntner M (2022) The new Australian leaf-curling orb-weaving spider genus Leviana (Araneae, Araneidae). Evolu- tionary Systematics 6(2): 103-133. https://doi.org/10.3897/evol- syst.6.83573 Framenau VW, Dupérré N, Blackledge TA, Vink CJ (2010) Systemat- ics of the new Australasian orb-weaving spider genus Backobourkia (Araneae: Araneidae: Araneinae). Arthropod Systematics & Phylog- eny 68: 79-111. Framenau VW, Baptista RLC, Oliveira FSM, Castanheira P de S (2021a) Taxonomic revision of the new spider genus Hortophora, the Aus- tralasian Garden Orb-weavers (Araneae, Araneidae). Evolutionary Systematics 5(2): 275-334. https://do1.org/10.3897/evolsyst.5.72474 Framenau VW, Vink CJ, Scharff N, Baptista RLC, Castanheira P de S (2021b) Review of the Australian and New Zealand orb-weaving spider genus Novakiella (Araneae, Araneidae). Zoosystematics and Evolution 97(2): 393-405. https://doi.org/10.3897/zse.97.67788 Framenau VW, Castanheira P de S, Vink CJ (2022) Taxonomy and systematics of the new Australo-Pacific orb-weaving spider genus Socca (Araneae: Araneidae). New Zealand Journal of Zoology 49: 263-334. https://doi.org/10.1080/03014223.2021.2014899 Joseph MM, Framenau VW (2012) Systematic review of a new orb-weaving spider genus (Araneae: Araneidae), with special ref- erence to the Australasian-Pacific and South-East Asian fauna. Zoo- logical Journal of the Linnean Society 166(2): 279-341. https://doi. org/10.1111/j.1096-3642.2012.00845.x 323 Kallal RJ, Hormiga G (2018) Systematics, phylogeny and biogeogra- phy of the Australasian leaf-curling orb-weaving spiders (Araneae: Araneidae: Zygiellinae), with a comparative analysis of retreat evolution. Zoological Journal of the Linnean Society 184(4): 1055— 1141. https://doi.org/10.1093/zoolinnean/zly014 Keyserling E (1886) Die Arachniden Australiens nach der Natur bes- chrieben und abgebildet. Bauer & Raspe, Nurnberg. Levi HW (1992)TheAmerican species of the orb-weaver genus Carepalx- is and the new genus Rubrepeira (Araneae: Araneidae). Psyche, Cambridge 98(2-3): 251-264. https://doi.org/10.1155/1991/26493 Rack G (1961) Die Entomologischen Sammlungen des Zoologischen Staatsinstituts und Zoologischen Museums Hamburg. 2. Teil, Che- licerata 11: Araneae. Mitteilungen aus dem Hamburgischen Zoolo- gischen Museum und Institut 59: 1-60. Rainbow WJ (1911) A census of Australian Araneidae. Records of the Australian Museum 9(2): 107-319. https://doi.org/10.3853 /}.0067-1975.9.1911.928 Scharff N, Coddington JA (1997) A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae). Zoo- logical Journal of the Linnean Society 120(4): 355—434. https://doi. org/10.1111/j.1096-3642.1997.tb01281.x Scharff N, Coddington JA, Blackledge TA, Agnarsson I, Framenau VW, Sziits T, Hayashi CY, Dimitrov D (2020) Phylogeny of the orb- weaving spider family Araneidae (Araneae: Araneoidea). Cladistics 36(1): 1-21. https://doi.org/10.1111/cla. 12382 Simon E (1908) Araneae, 1" partie. In: Die Fauna Stidwest-Australiens. Ergebnisse der Hamburger stidwest-australischen Forschungsreise 1905 (ed. W. Michaelsen, R. Hartmeyer). Verlag von Gustav Fisch- er, Jena, 359-446. World Spider Catalog (2023) World Spider Catalog Version 23.5. [Available from:] http://wsc.nmbe.ch [21 April 2023] Yin CM, Wang JF, Zhu MS, Xie LP, Peng XJ, Bao YH (1997) Fauna Si- nica: Arachnida: Araneae: Araneidae. Science Press, Beijing, 460 pp. zse.pensoft.net