A large brush-footed trapdoor spider (Mygalomorphae: Barychelidae) from the Miocene of Australia Author McCurry, Matthew R. Australian Museum Research Institute, Sydney, NSW 2010, Australia & Earth & Sustainability Science Research Centre, School of Biological, Earth and Environmental Sciences (BEES), University of New South Wales, Kensington, NSW 2052, Australia & Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA matthew.mccurry@australian.museum Author Frese, Michael Australian Museum Research Institute, Sydney, NSW 2010, Australia & Commonwealth Scientific and Industrial Research Organisation, Health and Biosecurity, Black Mountain, ACT 2601, Australia & Faculty of Science and Technology, University of Canberra, Bruce, ACT 2601, Australia Author Raven, Robert Biodiversity and Geosciences Program, Queensland Museum, South Brisbane, QLD 4101, Australia text Zoological Journal of the Linnean Society 2024 2023-09-15 200 4 1026 1033 http://dx.doi.org/10.1093/zoolinnean/zlad100 journal article 10.1093/zoolinnean/zlad100 0024-4082 11240583 Genus Megamonodontium gen. nov. Remarks: The similarity in the length of the legs ( Fig. 2 ) suggest that the genus belongs to the Tuberculotae sensu Raven (1985) and amongst them, shows the strongest affinities with the Barychelidae , which also have truncated-looking tarsi ( Fig. 3A ; see Raven 1994 : Tarsal Extremity, p. 313 and fig. 12A–Q). The fluted ridges on setae on the legs are also consistent with those in the Barychelidae (see Raven 1994 : figs 21F, 22A). Figure 1. Part (A) and counterpart (B) of Megamonodontium mccluskyi ( AM F.145559 ). Figure 2. Annotated composite line drawing of AM F.145559 , created using the part and counterpart of the fossil. The parts studied have cleaved through the carapace and abdomen, meaning that, as noted, the chelicerae, eyes, fovea, sternum and spinnerets are unknown, as are the distal ends of the first, third and fourth legs on the counterpart. Much of the specimen shows the body wall from the inside of the specimen; only in some areas are external details of the body evident. The specimen is compressed but shows some three-dimensional relief. Almost all of what we know about this spider derives from the morphology of the legs. However, of the 185 characters used by Raven (1985) , 42 are based upon the legs including the claws and spines, excluding the male palp and coupling spurs. The fossil surface visible seems to be that immediately below the cuticle, with only setal scars evident ( Fig. 3F, G ). Despite the limited number of characters, we can still place the specimen reliably within the brush-footed trapdoor spiders. Raven (1985) posited that the mygalomorphs form two groups, based upon the relative sizes of their legs. The Atypoidea and Rastelloidina (Fornicephale) all have noticeably weaker front (I and II) legs than rear (III and IV) legs ( Fig. 4A–C ); also, in Atypoidea and Rastelloidina, the spination of the distal segments (tibia to tarsi) of legs I and II is very strong; the spines on the third leg are the strongest and those on the fourth leg are the weakest ventrally. Although a number of other characters, such as the presence of digging spines (rastellum) on the chelicerae, the reduction of the eye tubercle, the absence of dense hair (scopula) on the legs, and the steeply elevated head region or caput are not discernible on this specimen, we assume that they are present, because they are strongly correlated with the spination characters ( Raven 1985 ). Figure 3. Scanning electron micrographs of Megamonodontium mccluskyi . A, B, pedipalp with tarsal claw. C, D, seta located on the pedipalp. E, F, internal structure of abdominal setae attachments. G, setal attachments on the internal cuticle surface, showing impressions of setae projecting on the outside of the cuticle. H, close-up of a seta base. Scale bars: 250 µm (A, E); 100 µm (B, G); 50 µm (C, F); 25 µm (H); 5 µm (D). Raven’s second group are the Tuberculotae, which are characterized by eyes raised on a common tubercle, a lower head region, and often hirsute distal leg segments. The Tuberculotae have legs of similar thickness ( Fig. 4D–F ) and weak, if any spines on the legs apart from the third and fourth leg, and it is to this group that this species clearly belongs. Further correlated with those legs are the reduction in size of the third or middle claw on the legs and the development of a pair of pads around the claws, claw tufts, as found in the tarantulas and the brush-footed trapdoor spiders, Barychelidae ( Raven 1994 ) . The inclusion of this species in the brush-footed trapdoor spiders is challenged by the apparent absence of claw tufts on the tarsi. The only fossil relative of brush-footed trapdoor spiders ( Psalistops Simon, 1889 , now in the Theraphosidae ; Mori and Bertani 2020 ) had its affinities recognized because a preservation in amber ( Wunderlich 1988 ) made it possible to observe the tufts. Within the Tuberculotae, the Crassitarsae were so named for those with hirsute matts (scopula) of setae on the ventral metatarsi and tarsi, at least of legs I and II ( Raven 1985 ). The best developed of those scopulae are found in the tarantulas and the brush-footed trapdoor spiders, Barychelidae ( Raven 1985 , 1994 ). In those families, the relative lengths of the leg segments are a taxonomically useful character: as in most other spiders, the patella of the first and second leg of females is usually the shortest segment, but in many brush-footed trapdoor spiders, the patellae of legs I and II are longer than their respective tibiae ( Raven 1994 ). This contrasts with the usually short patellae in females of their putative sister group, the tarantulas ( Raven 1985: 311 ; 2005: 311 ). Holotype : AM F.145559 , part ( Fig. 1A ) and counterpart ( Fig. 1B ) of a single compressed specimen, registered and housed in the palaeontology collection of the Australian Museum, Sydney, NSW, Australia (LSID Urn:lsid:zoobank.org:act: 51339D5C-590D-424B-B3FD-0CCF02126C44 ). Etymology : The genus name alludes to the nearest living relatives, tiny litter-dwelling brush-footed trapdoor spiders of the genus Monodontium . Locality: The holotype derives from McGraths Flat, a fossil site that is located ~ 25 km north-east of Gulgong, MSW, Australia . The fossil is preserved in a finely bedded goethite matrix with low levels of silica present, which has been dated to the Miocene (11–16 Mya), based on pollen and spores ( McCurry et al. 2022 ). Diagnosis: Unlike most brush-footed trapdoor spiders, Megamonodontium has females with patella I shorter than tibia I ( Fig. 1 ) and teeth on both the palpal claw ( Fig. 3A, B ) and the outer face of the paired tarsal claws of females. Megamonodontium differs from Tungari kenwayae Raven, 1994 in that the first leg is longer than the second and from all species of Zophorame in that its legs are robust, that it lacks well-developed scopulae, and that its head and abdomen are less slender. Megamonodontium differs from its putative sister genus, Monodontium , in having the carapace shape asymmetrical around the midpoint, with the widest point in the posterior third; Monodontium has the carapace asymmetrical around the midpoint ( Raven 2008 : fig. 8A, C); it is also about five times the size of known Monodontium (10 vs. 2 mm ). Remarks: Megamonodontium does not have visible scopulae, a condition that is considered one of the key features of the Crassitarsae. Putatively, the most plesiomorphic genus in the Crassitarsae is the barychelid genus Monodontium Kulczyski, 1908 revised by Raven (2008) . Although a brush-footed trapdoor spider, Monodontium has several highly plesiomorphic characters, including hardly developed scopulae and claw tufts. Amongst its species, most females have relatively long patellae I and II, but Monodontium sarawak Raven, 2008 has a female with short patellae I and II ( Raven 2008 ). Equally, in Monodontium , not only the males, but also the females have biserially dentate, paired claws. Megamonodontium shares this trait, having teeth on the outer face of the paired tarsal claws of females ( Fig. 3A, B ). In most species of Tungari Raven, 1994 and the New Caledonian Barycheloides alluviophilus Raven, 1994 and Barycheloides chiropterus Raven, 1994 , females have teeth on the paired claws of the first leg but none on the palpal claw. In species from the Cape York Peninsular in Northern Australia , Tungari kenayae Raven, 1994 and all species of Zophorame Raven, 1990 , females also have teeth on the palpal claw and on the paired claws of the first leg, but Megamonodontium differs from these species in many other ways, including the length and proportions of the legs, the shape of the cephalothorax and the shape of the abdomen. In other barychelids, the paired claws of females are either edentate or have only few teeth and then medially, not in a row on each face of the claw. Raven (1985) posited that this condition derived from the sister group of the Theraphosidae and Barychelidae (plus Paratropididae ), the Nemesiidae (now split into Anamidae , Bemmeridae and Nemesiidae ). In the Nemesiidae s.l. , both males and females of all but one genus ( Spiroctenus ) have the paired claws biserially dentate. In the tarantulas, males of the plesiomorphic Ischnocolinae are like those of most brush-footed trapdoor spiders and have biserially dentate paired claws, whereas females have little or no teeth. The most recent phylogeny of spiders that includes both Barychelidae and Theraphosidae is that of Goloboff-Szumik and Ríos-Tamayo (2022) , based upon the work of Mori and Bertani (2020) . This study places the brush-footed trapdoor spiders as the sister group of the tarantulas but widely disperses other taxa with biserial dentition of the paired tarsal claws, intermixing them with small-bodied or monoserially dentate taxa. Older studies have yielded similar results ( Wheeler et al. 2017 , Opatova et al. 2020 ). Within the Tuberculotae ( Fig. 3D–F ), the Idiopidae and the Nemesiidae s.l. can be eliminated by the highly reduced leg spination in Megamonodontium , in addition to the dentition of the palpal claw. Hence, by a process of elimination, using only the leg characters in the species, we conclude that this species is a close relative of the brush-footed trapdoor spider Monodontium , a tropical genus of dipterocarp rainforest dwellers. Figure 4. A–C, Fornicephalae (females) showing reduced length and diameter of legs I and II, as depicted by Raven (1985) : A, Conothele Thorell, 1878 (Eromanga, Queensland), Halonoproctidae ; B, Missulena bradleyi Rainbow, 1914 (Brisbane, Queensland), Actinopodidae ; C, Migas plomleyi Raven and Churchill, 1989 (Launceston, Tasmania), Migidae . D–F, Tuberculotae (females) showing similar length and diameter of legs I, II–IV: D , Idiommata Ausserer, 1971 (Gladstone, Queensland), Barychelidae ; E, Cethegus Thorell, 1881 (Normanton, Queensland), Euagridae ; F, Hadronyche formidabilis Rainbow, 1914 (Bunya Mountains, Queensland), Atracidae . Type species Megamonodontium mccluskyi sp. nov. Etymology : The species is named after Simon McClusky, who found the specimen. Diagnosis: The diagnosis is as for the genus. Description: Almost complete part and counterpart of an adult female spider. Moderately large spider (carapace length, ~ 10 mm ), with stout legs. The femora of legs I and II are thicker than those of III and IV. Palpal claw with three teeth distally on the claw ( Fig. 3A, B ); pedal claws (at least on leg III) with at least three teeth on one face of paired tarsal claws, presumably on both faces. Several spines on the prolateral face of the palpal tarsus ( Fig. 3A, B ) and at least one on prolateral tibia III; few spines present on legs; no spines on pedal tarsi. Setae with anastomosed ridges ( Fig. 3C, D ). Abdomen ovate, with pustules ( Fig. 3E, F ) across the dorsal and posterior surface; a pair of book lungs evident. Chelicerae, eyes, fovea, sternum, spinnerets and male unknown. Additional measurements are presented in Table 1. A paired set of book lungs can be observed within the anterior portion of the abdomen ( Figs 1 , 2 ). There is a light-yellow-coloured structure around the midline of the anterior portion of the cephalothorax. This structure appears in the approximate position of the epipharyngeal ganglion, but there are no morphological features present to confirm its identity ( Figs 1 , 2 ). However, another fossil spider has been found at McGraths Flat that provides further evidence for the preservation of neuronal tissues. Richardson et al. (2023) report a jumping spider (Aranaeae: Salticidae ) that contains ‘a putative neuropile’ composed of ‘semi-parallel bundles of mostly hollow tubes with a diameter of 0.5–1 μm that have the general appearance of axons’.