T A X O NOM I C A R T I C L E Integrative taxonomic revision of the Australian cave cricket Speleotettix Chopard, 1944 (Orthoptera: Rhaphidophoridae): New species, distribution and conservation implications Perry G. Beasley-Hall1 | Steven A. Trewick2 | Brock A. Hedges1 | Steven J. B. Cooper1,3 | Elizabeth H. Reed1 | Andrew D. Austin1 1School of Biological Sciences, University of Adelaide, Adelaide, South Australia, Australia 2Wildlife and Ecology Group, Massey University, Palmerston North, New Zealand 3Evolutionary Biology Unit, South Australian Museum, Adelaide, South Australia, Australia Correspondence P. G. Beasley-Hall, School of Biological Sciences, The University of Adelaide, Adelaide, South Australia, Australia. Email: perry.beasley-hall@adelaide.edu.au Funding information This work was funded by a fellowship awarded to PGBH through the National Taxonomy Research Grant Program by the Australian Government’s Department of Climate Change, Energy, the Environment and Water (no. 4-H3JJWE), with co-funding from the Australian Speleological Federation, South Australian Museum, Environment Institute and the University of Adelaide. Abstract Cave crickets, also called w�et�a, are an important component of subterranean realms globally, but the true diversity of the group is poorly known. Speleotettix Chopard, 1944 contains two species from southeast Australia, Speleotettix tindalei Chopard, 1944 and Speleotettix chopardi (Karny, 1935). However, the initial descrip- tion of the genus was poorly characterised, and its taxonomy has remained unclear since. Here, we take an integrative molecular and morphological approach to redescribe Speleotettix and describe three new species: Speleotettix aolae Beasley-Hall, sp. nov. from Victoria and Speleotettix binoomea Beasley-Hall, sp. nov. from New South Wales, both found in limestone caves, and Speleotettix palaga Beasley-Hall, sp. nov. from mineshafts in Victoria. To provide consistency in future work on the group, we also redescribe S. tindalei and treat S. chopardi as a nomen dubium. Finally, we transfer the threatened species Cavernotettix crag- giensis Richards, 1974, an island species from Tasmania’s Bass Strait, into the genus as Speleotettix craggiensis (Richards, 1974) comb. nov. In so doing, we increase the number of described rhaphidophorid species in Australia to 27 and significantly expand the distribution of Speleotettix, making it the most widespread of the Australian genera currently known. As all members of Speleotettix are short-range endemics at risk of decline, these findings have implications for their future con- servation management. K E YWO R D S Macropathinae, w�et�a INTRODUCTION The orthopteran family Rhaphidophoridae Walker, 1869 is a globally distributed group of insects commonly known as cave crickets or w�et�a. Despite their common name, species are found in a range of habitats unified by cool, humid microclimates and low light levels, including wet forest, mineshafts and animal burrows (Lavoie & Poulson 2007). The subfamily Macropathinae Karny, 1930 represents the entirety of the rhaphidophorid fauna in the Perry Beasley‐Hall: http://zoobank.org/urn:lsid:zoobank.org:author:D8C4DBEA-5C2D-4A07-AEF7-0842ABFB708C http://zoobank.org/urn:lsid:zoobank.org:pub:455CB2A7-8DC9-4C47-8559-E30AD90556EC Received: 22 July 2024 Revised: 17 January 2025 Accepted: 7 April 2025 DOI: 10.1111/aen.70011 This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes. © 2025 The Author(s). Austral Entomology published by John Wiley & Sons Australia, Ltd on behalf of Australian Entomological Society. Austral Entomology. 2025;64:e70011. wileyonlinelibrary.com/journal/aen 1 of 20 https://doi.org/10.1111/aen.70011 https://orcid.org/0000-0002-7360-7933 mailto:perry.beasley-hall@adelaide.edu.au http://zoobank.org/urn:lsid:zoobank.org:author:D8C4DBEA-5C2D-4A07-AEF7-0842ABFB708C http://zoobank.org/urn:lsid:zoobank.org:pub:455CB2A7-8DC9-4C47-8559-E30AD90556EC https://doi.org/10.1111/aen.70011 http://creativecommons.org/licenses/by-nc/4.0/ http://wileyonlinelibrary.com/journal/aen https://doi.org/10.1111/aen.70011 http://crossmark.crossref.org/dialog/?doi=10.1111%2Faen.70011&domain=pdf&date_stamp=2025-07-01 Southern Hemisphere except for Southeast Asia and Papua New Guinea. Recent work has dealt with the sys- tematics and taxonomy of New Zealand rhaphidophorids (Fitness et al. 2018; Hegg, Morgan-Richards, & Trewick 2019, 2022; Johns & Cook 2013; Trewick 1999), but no such research has been conducted on Australian species for over five decades (Richards 1974). Nine rhaphidophorid genera are described from Australia: Australotettix Richards, 1964; Cavernotettix Richards, 1966; Eburnocauda Beasley-Hall & Iannello, 2024; Micropathus Richards, 1964; Novotettix Richards, 1966; Pallidotettix Richards, 1968; Parvotettix Richards, 1968; Speleotettix Chopard, 1944; and Tasmanoplectron Richards, 1971. To date, morphological characters used by Richards correlate well with phylogenetic relationships and existing generic assignments hold up well under molecular scrutiny (Beasley-Hall et al. 2018, 2025). How- ever, Speleotettix is poorly known because it is the only genus not described by Richards, meaning its descriptions were based on a different set of morphological characters and therefore cannot easily be compared with the remainder of the Australian fauna. The genus presently contains two species, Speleotettix tindalei Chopard, 1944 from the Limestone Coast of South Australia and Speleo- tettix chopardi (Karny, 1935) from the Dandenong Ranges in Victoria. The descriptions of both species (and the genus itself) are inadequate and require revision, primar- ily because neither provide sufficient morphological infor- mation for confident identification of taxa. Several additional species putatively assigned to Spe- leotettix have remained undescribed for decades. Richards deposited specimens from caves in central New South Wales (NSW) into museum collections under the tag name Speleotettix profundus in the 1960s, but this taxon was never formally described. In a subsequent species checklist for Australia, Richards alluded to an undescribed member of Cavernotettix Richards, 1966 from the “central coastal ranges” et al. of NSW with no mention of S. profundus (Richards, 1987). The shared location and morphological similarity between the two genera suggest Richards was referring to a single taxon with an uncertain generic placement. Richards also referred to two other undescribed species of Speleotettix from lava tubes and woodland in “coastal southwest Victoria” (VIC) but pro- vided no further information on their morphology or exact locality (Richards 1987). A taxonomic revision of Speleotettix is necessary not only because of the above issues but because most, if not all, members of the genus are short-range endemics likely deserving of conservation listing. Robust, accurate descriptions and keys are necessary for identification and subsequent conservation management of species (Dubois 2003; Vogel Ely et al. 2017), and a formal scientific name is generally required for a taxon to be protected under Australian environmental legislation and the Inter- national Union for Conservation of Nature’s Red List of Threatened Species (IUCN Standards and Petitions Committee 2024; Murphy & van Leeuwen 2021). As such, here we undertake a complete revision of Speleotettix to lay the groundwork for future taxonomic and conserva- tion work on the group. We begin by redescribing the genus and the two existing species therein. We then revisit locations mentioned by Richards to house unde- scribed taxa and describe three new species, Speleotettix aolae Beasley-Hall, 2025 sp. nov., Speleotettix binoomea Beasley-Hall, 2025 sp. nov. and Speleotettix palaga Beasley-Hall, 2025 sp. nov. The generic identity of these taxa within Speleotettix was confirmed in a recent molecu- lar phylogeny by the authors (Beasley-Hall et al. 2025) (Figure 1). Finally, we redescribe Cavernotettix craggiensis Richards, 1974 as a member of Speleotettix following its recovery within the genus based on the aforementioned phylogenetic study (Figure 1). MATERIALS AND METHODS Taxon sampling and imaging Specimens were collected from the field and loaned from museum collections. Crickets were identified as Speleotettix as a first pass based on the coxae lacking anterolateral spines, the hind femora bearing an apical retrolateral spine only, and the male suranal plate being developed into two elongate processes. Identification was further confirmed using mitochondrial and nuclear molecular data presented in a sister study by Beasley-Hall et al. (2025). Samples were imaged using a Canon EOS 5DS R and a Canon MP-E65mm f/2.8 1-5� macro lens. Images were processed using Zerene Stacker v.1.04 with the PMax stacking method, Adobe Photoshop 2025 and Adobe Lightroom 2025. DNA sequencing and phylogenetic analysis In addition to justifying our descriptions with molecular data from Beasley-Hall et al. (2025), we also sequenced the cytochrome oxidase I (COI) gene from new material depos- ited at the Australian Museum from Junction and Fig Tree caves at Wombeyan, NSW (Figure 1). The identity of this population was previously uncertain as it was recovered as multifurcating (i.e., with a polytomy) by Beasley-Hall et al. (2025), albeit with a close relationship to populations at Jenolan and Abercrombie. No new material from Abercrombie was available to us in the present study. DNA extractions of fresh specimens from Wombeyan were performed using a Gentra Puregene Tissue Kit (QIAGEN), and we amplified COI using the primers LCO1490 + HCO2198 (Folmer et al. 1994). PCRs were per- formed using a standard hot-start protocol consisting of a preheat step at 95�C for 10 min, denaturation at 95�C for 45 s, annealing at 48�C for 45 s, extension at 72�C for 1 min, 72�C for 10 min and 25�C for 2 min 20 s. Dena- turation, annealing and extension steps were repeated for 2 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense 39 cycles. Product purification and DNA sequencing was outsourced to the Australian Genome Research Facility (Melbourne, Australia). COI sequences from the two Wombeyan samples were aligned against barcodes from Beasley-Hall et al. (2025) using MUSCLE v.3.8.425 (Edgar 2004) for a total of 678 bp. The dataset was parti- tioned by codon position. Phylogenetic analysis was con- ducted using IQTREE v.1.6.12 with 1000 ultrafast bootstrap replicates and substitution model selection per- formed by ModelFinder (MFP + MERGE option) (Hoang et al. 2018; Kalyaanamoorthy et al. 2017). We manually rooted the tree at the split between S. binoomea and all other members of the genus based on the topology recovered by Beasley-Hall et al. (2025). Species delimitation We conducted species delimitation analyses to support our morphological descriptions of new members of Speleotettix. Here we followed the unified species concept (USC) of De Queiroz (2007), which conceptualises species as “separately evolving metapopulation lineages” (or seg- ments thereof) supported by lines of evidence which may include reciprocal monophyly, fixed morphological differ- ences or reproductive isolation. In keeping with the USC and to avoid potentially over- or under-splitting lineages by relying on a single method in isolation (Luo et al. 2018), we used several different approaches to test our species hypothesis: (1) the distance-based Assemble Species by Automatic Partitioning method (ASAP) (https://bioinfo. mnhn.fr/abi/public/asap/asapweb.html; Puillandre, Brouillet, & Achaz 2021); (2) the distance and tree-based Species Delimitation plugin (SDP) (v.1.4.5; Masters, Fan, & Ross 2011) in Geneious Prime v.2024.0.3 (http://geneious. com); and (3) the tree-based Bayesian implementation of the Poisson tree process (bPTP) (https://species.h-its.org/ ptp/; Zhang et al. 2013). The ASAP method identifies barcode gaps without the need for a user-specified prior on intraspecific divergence and can rank alternative partitioning schemes. We ran the analysis using Kimura 2-parameter distances and a probability threshold of 0.01. We next used SDP, which relies on a single gene tree to calculate average pairwise intraspecific and interspecific genetic differences between putative species. The SDP plugin also calculates the probability of reciprocal monophyly stemming from evolutionary lineage separa- tion vs. a random outcome of the coalescent process using Rosenberg’s PAB, with a value of <0.05 indicating strong evidence for delimitation between species (Rosenberg 2007). Finally, we used bPTP, a tree-based method that relies on the number of substitutions along branches to infer species boundaries. We ran bPTP for 100 000 MCMC generations using a thinning of 100 and a 10% burn-in. We were not able to include F I G U R E 1 (a) Phylogenetic relationships among members of Speleotettix adapted from Beasley-Hall et al. (2025). Cavernotettix craggiensis is assigned to the genus in the present paper. Samples from caves at Abercrombie and Wombeyan were putatively assigned to sp. nov. 3 by those authors, but their position remains unclear given polytomies in the relevant clade (red question mark). (b) Locations of populations shown in (a) as well as Speleotettix chopardi, the second described species in the genus. REVISION OF THE CAVE CRICKET Speleotettix 3 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense https://bioinfo.mnhn.fr/abi/public/asap/asapweb.html https://bioinfo.mnhn.fr/abi/public/asap/asapweb.html http://geneious.com http://geneious.com https://species.h-its.org/ptp/ https://species.h-its.org/ptp/ S. chopardi in these analyses due to its status as a nomen dubium (see Taxonomy section below). Abbreviations and terminology Collection abbreviations iNaturalist = Available from iNaturalist.org (as of 21 May 2024). ANIC = Australian National Insect Collection, CSIRO, Canberra AM = Australian Museum, Sydney QM = Queensland Museum MNHN = Muséum National d’Histoire Naturelle, Paris Locations NSW = New South Wales SA = South Australia TAS = Tasmania VIC = Victoria We use ‘single quotation marks’ to indicate unofficial taxonomic or location names, as for S. profundus, and “double quotation marks” to indicate exact quotes from published texts. Morphological terminology The terminology for rhaphidophorid morphology is inconsistent in the literature. Here, we follow the conven- tions of Richards (1964a) and more recent studies of Macropathinae (Fitness et al. 2015; Hegg et al. 2022) in referring to terminalia and the spination of the legs. Api- cal spines are positioned at the distal end of a leg seg- ment, and subapical spines are positioned between a pair of apical spines, the latter only occurring on the hind tibia (Figure 2). Linear spines occur in rows along a leg segment and may occur on the dorsal (also superior) or ventral (also inferior) surface of that segment. Both apical and lin- ear spines may face closer to the head (prolateral) or the posterior of the body (retrolateral). Apical and linear spines are sometimes referred to as immovable spines and movable spurs, respectively, particularly when referring to the Northern Hemisphere Rhaphidophoridae (Jiao et al. 2008; Qin, Liu, & Li 2016; Zhou & Yang 2022). Further complicating matters, Richards sometimes confused her descriptions of spine placement and would refer to structures directed prolat- erally as being retrolateral, and vice versa (e.g., Richards 1964b, 1966b). This phrasing is particularly confusing for projections on the coxae which, unlike api- cal or linear spines of other leg segments, do not clearly face in either direction yet are helpful in distinguishing Speleotettix from related genera like Cavernotettix. We do not consider Richards’ terms appropriate for the coxae and instead follow Hegg et al. (2022) in referring to forward-facing processes as anterolateral spines and downward-facing processes as ventrally directed. RESULTS Species delimitation We used the molecular phylogeny of Beasley-Hall et al. (2025) (Figure 1) to construct an initial species hypothesis F I G U R E 2 Apical and subapical spines on the femora and tibiae can be used to distinguish rhaphidophorid genera. Pro = prolateral (p); ret = retrolateral (r); sup = superior surface; inf = inferior surface. Cross-sections, denoted by opaque boxes, are from a distal perspective. Diagram after Fitness et al. (2015). 4 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense http://iNaturalist.org for members of Speleotettix. We assessed this hypothesis using four different species delimitation methods in the present study (Figure 3). As COI was una- vailable for S. craggiensis comb. nov., it could not be included in our analyses. However, the species was confi- dently recovered within Speleotettix by Beasley-Hall et al. (2025) based on 12S rRNA data, and it is clearly separated from other members of the genus by its morphology, justifying its redescription as a new combination below. Similarly, S. chopardi was unavailable for examination given its status as a nomen dubium, making a confident identification impossible in our view (see Taxonomy section below). The division of Speleotettix into five separate molecu- lar lineages (not including S. craggiensis comb. nov. or chopardi) was supported by all species delimitation methods used here (Figure 3). Specimens of a new spe- cies at Wee Jasper were too immature for morphological examination but the population was confidently delim- ited from other members of Speleotettix based on molec- ular data (Figure 3). The distance-based ASAP and SDP methods separated lineages based on an interspecific genetic distance of 2.3%–4.4%, corresponding to the low- est ASAP score of 2.5 and a Rosenberg’s PAB of <0.05 for every lineage (Figure 3; Table S1). The tree-based bPTP method estimated between 3 and 15 species in our COI phylogeny, with the five-lineage scheme being the highest-scoring in both the maximum likelihood and Bayesian solutions. Support values for the delimitation of each species by bPTP ranged between 0.324 (S. tindalei) and 0.758 (Wee Jasper sp. nov.). Beasley-Hall et al. (2025) recovered two populations of cave crickets from Wombeyan and Abercrombie Karst Conservation Reserves as closely related to those at Jenolan (Figure 2), but their position was complicated by a polytomy in each case. Here, we examined new material from Wombeyan and generated additional COI data to confidently determine the identity of the population. Relationships among other Speleotettix inferred by Beasley-Hall et al. (2025) are otherwise robust, so we integrated this new data alongside existing COI sequences in a maximum likelihood phylogeny. We recovered the Wombeyan population as a member of S. binoomea sp. nov. with high node support (Figure 3) and confirmed its identity through morphological exami- nation. Material from Abercrombie was unavailable for examination in the present study but it likely also repre- sents a subpopulation of S. binoomea sp. nov. In contrast, an undescribed population from Wee Jasper (Figure 1) is differentiated from S. binoomea sp. nov. by over 7% pair- wise COI genetic distance, suggesting it is a new species in its own right rather than a subpopulation of S. binoomea (Figure 3; Table S1). TAXONOMY Infraorder Tettigoniidea Kevan, 1982. Superfamily Rhaphidophoroidea Walker, 1869. Family Rhaphidophoridae Walker, 1869. Subfamily Macropathinae Karny, 1930. Tribe Macropathini Karny, 1930. Genus Speleotettix Chopard, 1944. Type species: Speleotettix tindalei Chopard, 1944 by origi- nal designation. Included species: Speleotettix aolae Beasley-Hall sp. nov.; Speleotettix binoomea Beasley-Hall sp. nov.; Speleotettix chopardi (Karny, 1935); Speleotettix craggiensis (Richards, 1974) comb. nov.; Speleotettix palaga Beasley-Hall sp. nov.; Speleotettix tindalei Chopard, 1944. F I G U R E 3 Relationships among Speleotettix species based on COI, with new material incorporated from Wombeyan (NSW) in black. The result of four species delimitation methods is shown to the right. Morph. = morphology. Speleotettix craggiensis comb. nov. was excluded from distance and tree-based analyses due to a lack of COI data but is clearly delimited by morphology. Speleotettix chopardi, shown in Figure 1, was not sampled here because its description is too vague to facilitate species identification. REVISION OF THE CAVE CRICKET Speleotettix 5 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense Diagnosis Speleotettix comprises medium-sized crickets with an adult body length—that is, from the top of the fastigium to the distal margin of the suranal plate—of up to 19 mm and a distribution spanning the Bass Strait of TAS and southeast mainland Australia (SA, VIC and NSW) (Figure 1). The basic body colouration is beige with brown patterning. Unlike many other Australian Rhaphidophori- dae, the genus is identified by a combination of morpho- logical characters rather than a single trait. Speleotettix lacks spination on the coxae, also shared by Novotettix Richards, 1966 and Pallidotettix Richards, 1968 but differ- entiated by a deeply emarginate distal margin of the male suranal plate and fewer dorsal linear spines on the hind tarsi. Speleotettix can also be differentiated by the comparatively weak teeth on the female ovipositor, which are more well-pronounced and may be reflexed proximally in Novotettix and Pallidotettix. Redescription Head Light to mid brown patterning on vertex and frons, span- ning median ocellus. Eyes black. Fastigium higher than long, indented medianly and divided into two laterally compressed tubercles. Antennae long, scapes almost touching at their bases. Body Basic body colour in life pale beige to ochreous with light to mid brown patterning. Body moderately clothed with short, pale brown setae. A median dorsal line present, beginning at the fastigium and running down the length of the body and becoming less visible beyond the tho- racic nota. Two faint, glabrous patches resembling eye- spots on either side of the pronotum, which may appear murky or shadow-like compared with surrounding pat- terning, where the internal cryptopleura attaches to the underside of the plate. Prominent tubercles may be pre- sent at the distal margin of abdominal tergites, as in Spe- leotettix palaga sp. nov. Legs Legs light to mid brown and interspersed with patches of beige on hind femora. Coxae unarmed and lacking both anterolateral spines and ventrally directed processes. Api- cal spines constant in number among species, except for hind femora (see below). Fore and middle femora bearing one prolateral and one retrolateral spine at apex and with- out ventral linear spines. Fore and mid tibiae bearing both apical and dorsal linear spines: apex of tibiae with four spines, one pair directed prolaterally and retrolaterally on superior surface, other prolaterally and retrolaterally on inferior; dorsal linear spine counts on tibiae vary among species. Fore and mid tarsi unarmed. Apex of hind femora usually with a single retrolateral spine only, but variably bearing a small prolateral spine in some species. Four api- cal spines on hind tibiae, as on fore and mid tibiae, but supplemented by two pairs of subapical spines between each primary pair. Apical spines present in a single pair each on first and second hind tarsal segments. Third and fourth hind tarsal segments unarmed. Male terminalia Male suranal plate deeply emarginate such that the pos- terior margin resembles an upside-down V, forming two triangular lobes. Subgenital plate trilobed, lateral lobes bearing styli; median lobe generally longer than lateral lobes. Sternite IX, which precedes the subgenital plate, bearing a medium-sized median tubercle; margin between the two structures often poorly visible. Female terminalia Female suranal plate broadly oval-shaped, concave laterally and straight to moderately emarginate at the dis- tal margin. Female subgenital plate trilobed; lobes wide, median lobe generally longer than lateral lobes. Oviposi- tor ranging from blue-grey or beige (juveniles) to brick red (adults) in life, often becoming translucent amber fol- lowing ethanol preservation. Ovipositor with six to eight teeth ranging from very weakly to moderately produced at distal margin; vertical ridges often present on the median surface leading to each tooth, which may be lack- ing for the most distal and/or proximal. Dorsal valve and median surface of ovipositor unarmed. Remarks Speleotettix is the sister genus to Cavernotettix (Beasley- Hall et al., 2025; Figure 1). The two genera have similar morphologies and are difficult to distinguish in photo- graphs alone, particularly if the spines (or lack thereof) on the fore coxae and/or the male suranal plate is not visible. Identification is further complicated by the fact that the genera are reportedly sympatric east of the Melbourne metropolitan area (Mesa 1970), but this has not been con- firmed by the authors. Speleotettix comprises a northern and southern clade: one in NSW (S. binoomea sp. nov. and Wee Jasper sp. nov.) and the other in SA, VIC and TAS (S. tindalei, S. aolae sp. nov., S. craggiensis comb. nov. and S. palaga sp. nov.). Etymology Not explained by Chopard. From the Latin sp�el�e �o = cave and Ancient Greek téttix = cicada, also used to refer to crickets and grasshoppers. The genus name is masculine. 6 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense Distribution SA, VIC, NSW and the Bass Strait of TAS (Figure 1). All spe- cies of Speleotettix occur in aphotic habitats or in close proximity to them, presumably observed when foraging. In SA, far west VIC and NSW, Speleotettix occurs in the twi- light zone of limestone and marble caves. In the Central Victorian Uplands, S. palaga is known from mineshafts. In TAS, S. craggiensis has been found under boulders and in animal burrows. Key to species of Speleotettix 1. Female suranal plate with a distinct median notch at the distal margin (Figure 5d). Lobes of female subgeni- tal plate subtriangular, lateral lobes longer than median lobe and plate uniformly sclerotised (Figure 5e). Gen- eral body colour mid brown and beige to ochreous (Figure 6). ………………………….binoomea sp. nov. Female suranal plate straight or only weakly emargin- ate at the distal margin (Figure 9d). Median lobe of female subgenital plate longer than lateral lobes (Figure 9e). General body colour mid brown and beige (Figure 10). …………………………………………….2 2. Female ovipositor with moderately produced teeth. Teeth most easily counted by their dentate disruption of the dorsal margin (Figure 4g). …………………...…3 Female ovipositor with weakly produced teeth, dorsal margin almost flat. Teeth most easily counted by ridges on the median surface of the ovipositor (Figure 9g). …………………………………………….4 3. Lateral lobes of female subgenital plate subrectangular with corners rounded distolaterally; distal margin of lat- eral lobes sclerotised and mid to dark brown, coloura- tion sometimes extending to the median surface of lobes (Figure 8e). Distal margin of female suranal plate emarginate (Figure 9d). Ovipositor with six teeth on the dorsal margin (Figure 9f).…...………….palaga sp. nov. Lateral lobes of female subgenital plate rectangular, distal margin of plate truncate. Distal margin of female suranal plate straight (Figure 8d). Ovipositor with eight teeth on the dorsal margin (Figure 8f). ………………………………...craggiensis comb. nov. 4. Lateral lobes of female subgenital plate not produced, flat; distal margin of lobes sclerotised and mid brown (Figure 9e). Distal margin of female suranal plate weakly emarginate (Figure 11d).………………………...tindalei Lateral lobes of female subgenital plate rounded; median surface margin of lateral lobes sclerotised and mid brown (Figure 4e). Distal margin of female suranal plate straight to weakly emarginate (Figure 4d). …………………………………………...aolae sp. nov. Speleotettix aolae Beasley-Hall, sp. nov. (Figure 4) http://zoobank.org/urn:lsid:act:10E6D414-5308-472C- 896F-70815A94C0D4 Speleotettix sp. nov. 1 (Beasley-Hall et al., 2025) Material examined Holotype Adult ♂, Cave BR5 (Big Cave), Portland, VIC, �38.335746, 141.508171, coll. S. Iannello, 18 March 2023, ANIC 14-009156. Paratype ?Subadult ♀, same locality and collection information, ANIC 14-009157. Other material 2 ♂, 2 ♀, same locality and collection information, ANIC 14-009158–14-009161. Jackass Fern Gully Picnic Area near Fitzroy River, VIC (photograph only), �38.074639, 141.425361, iNaturalist 119041985. Diagnosis Speleotettix aolae can be differentiated from other members of Speleotettix by the female terminalia. The species has a suranal plate with a straight to weakly emarginate distal margin and the subgenital plate bears lateral lobes which are rounded, shorter than the median lobe and darker than the surrounding plate due to sclerotisation. The ovipositor has eight weakly produced teeth on the dorsal margin. Description Measurements Holotype body length 14 mm, hind tibia 20 mm. ♀ para- type 11 mm, hind tibia 21 mm, ovipositor 11 mm. Head Light brown banding on vertex and frons of head forming a rough M shape, extending to a horizontal band that spans the fastigium. REVISION OF THE CAVE CRICKET Speleotettix 7 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense http://zoobank.org/urn:lsid:act:10E6D414-5308-472C-896F-70815A94C0D4 http://zoobank.org/urn:lsid:act:10E6D414-5308-472C-896F-70815A94C0D4 https://www.inaturalist.org/observations/119041985 Body Body generally mid brown and beige, with brown mot- tling covering thoracic nota and abdominal tergites and interrupted by beige patches. Small, well-defined spots generally concentrated on either side of the dorsal median line and at the distal margin of nota and tergites, with larger patches occurring laterally. Patterning mir- rored on either side of the dorsal median line and con- tinuing in this manner down the body. Legs Legs generally light brown and interspersed with beige patches, with the hind femora also bearing beige stria- tions laterally. Hind tibia with equidistant beige spots dor- sally, typically coinciding with pairs of larger dorsal linear spines. Fore and mid femur without ventral linear spines. Inferior surface of fore and mid tibia armed with two rows bearing an average of three dorsal linear spines each, one prolateral and one retrolateral. Ventral surface of hind femur bearing an average of 56 (min. 45, max. 62) prolateral and 41 (39–44) retrolateral linear spines in males and 9 (7–11) and 10 (9–11), respectively, in females. Dorsal surface of hind tibia with an average of 51 (46–58) prolateral and 48 (43–52) retrolateral linear spines in both sexes. First segment of hind tarsus with an average of 3 (2–5) dorsal linear spines prolaterally and 3 (1–4) retrolaterally. Second tarsal segment bearing an average of 2 dorsal linear spines, 1 (1–2) each prolaterally and retrolaterally. Male terminalia Suranal plate predominantly beige and light brown, with dark brown at the lateral margins; plate sparsely clothed with short setae. Distal margin of plate thickly clothed with longer setae and strongly emargin- ate. Subgenital plate light brown and sparsely clothed with short brown setae; a rounded, poorly pronounced median lobe at the distal margin, longer than lateral lobes. Female terminalia Suranal plate beige medianly and mid to dark brown lat- erally. Proximal margin of plate rounded, distal margin straight to weakly emarginate. Entire surface of plate sparsely covered in short, brown setae and with longer setae densely concentrated at the distal margin. Subge- nital plate distorted, but appearing sclerotised and F I G U R E 4 Speleotettix aolae Beasley-Hall, sp. nov. Head (a); dorsal view of terminalia (b ♂, d ♀); ventral view of terminalia (c ♂, e ♀); lateral view of ovipositor (f, g); dorsal view of hind tibia and tarsus (h). Scale bar = 1 mm. 8 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense bearing three beige lobes; lateral lobes rounded, median lobe broad with a triangular apex and longer than lat- eral lobes. Ovipositor bearing eight weakly produced teeth. Remarks The female specimens of S. aolae examined here did not exceed 11 mm in body size and have very poorly pro- duced teeth on the dorsal valve of the ovipositor (Figure 4f,g), suggesting they may be subadults. We have nonetheless included the shape of the teeth in our key to the genus as the species is sister to S. tindalei, in which this trait is present in adults. We also note the subgenital plate in the female paratype is distorted (Figure 4e); its general shape and degree of sclerotisation can still be gauged, but it is likely the plate is not cross-shaped in fresher material. Etymology Dedicated to Dr Aola Richards (1927–2021), who described the majority of the Australian and New Zealand Rhaphido- phoridae. It is a feminine noun in the genitive case. Distribution Limestone caves in the Portland area, VIC, Australia. Speleotettix binoomea Beasley-Hall, sp. nov. (Figure 5) http://zoobank.org/urn:lsid:act:0F07C512-3B3D-423D- B004-1DC2E7956C17 Cavernotettix sp. (Eberhard, 2014). Speleotettix sp. nov. 3 (Beasley-Hall et al., 2025) F I G U R E 5 Speleotettix binoomea Beasley-Hall, sp. nov. Head (a); dorsal view of terminalia (b ♂, d ♀); ventral view of male and female terminalia (c ♂, e ♀); lateral view of ovipositor (f,g); dorsal view of hind tibia (h). Scale bar = 1 mm. REVISION OF THE CAVE CRICKET Speleotettix 9 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense http://zoobank.org/urn:lsid:act:0F07C512-3B3D-423D-B004-1DC2E7956C17 http://zoobank.org/urn:lsid:act:0F07C512-3B3D-423D-B004-1DC2E7956C17 Material examined Holotype Adult ♂, entry to Orient Cave, Jenolan Karst Conservation Reserve, NSW, �33.821389, 150.022194, coll. A. Musser and M. Collins, 9 June 2023, AM K.621932. Paratypes Adult ♀, Binoomea Cut, Jenolan Karst Conservation Reserve, NSW, �33.821, 150.022, coll. A. Musser, 10 June 2023, AM K.621931. Other material: ♂, same locality and collection information as holotype, AM K.621930. 3 ♀ (AM K.627706, K.627707, K.627710), 2 ♂ (AM K.627708, K.627709), Hennings Cave J76 entrance chamber, Jenolan Karst Conservation Reserve, NSW, �33.798833, 150.017611, coll. H. M. Smith, A. Musser and T. Willmore, 13 June 2023. 1 ♂, Hennings Cave J76 entrance chamber, Jenolan Karst Conservation Reserve, NSW, �33.821694, 150.022194, coll. A. Musser, H. M. Smith and T. Willmore, 13 June 2019, AM K.386399. 1 ♂, entry to Temple of Baal Cave (photograph only), Jenolan Karst Conservation Reserve, NSW, �33.818944, 150.020806, iNaturalist 184652560. 2 ♀ (AM K.638921, K.638922), 2 ♂ (AM K.638925, K.638926), Junction Cave, Wombeyan Karst Conservation Reserve, NSW, coll. H. M. Smith, A. Musser and T. Willmore, 23–24 March 2024; GenBank no. for AM K.638921 PV461561. 1 ♀ (AM K.638923), 1 ♂ (AM K.638924), Fig Tree Cave, Wombeyan Karst Conservation Reserve, NSW, same collection information as previous; GenBank no. for AM K.638923 PV461562. 1 ♂, 1 ♀, Junction Cave, Wombeyan Karst Conservation Reserve, NSW, coll. F. Stone and T. Matts, 19 February 1997, QM (unregistered material, loan no. ENT22.19). Sex not recorded, Fig Tree Cave, Wombeyan, NSW, coll. A. M. Richards, 2 September 1961, ANIC 14-009132 (molecular data only). Sex not recorded, Abercrombie Caves, NSW, coll. E. Slater, 29 May 1960, ANIC 14-009131 (molecular data only). Diagnosis Speleotettix binoomea can be differentiated from other members of Speleotettix by its body colour and the female terminalia. The species is generally beige or ochreous, the latter particularly prominent in males from Jenolan but also evident at Wombeyan (Figure 10). The suranal plate is notched at the distal margin and the subgenital plate’s lateral lobes are longer than the median lobe. The ovipositor has eight moderately produced teeth on the dorsal margin. Description Measurements Holotype body length 16 mm, hind tibia 25 mm. ♀ para- type 11 mm, hind tibia 21 mm, ovipositor 10 mm. Head Delicate, light brown patterning on vertex and frons of head resembling leaf veins, extending below to a more concentrated section of brown spanning the fastigium. Tubercles of fastigium strongly produced. Body Median dorsal line on thoracic nota thin and often not extending past metanotum. Body generally mid brown and ochre—the latter more pronounced in males—with brown mottling on thoracic nota and abdominal tergites and interrupted by beige or ochreous patches. Small, well-defined spots generally concentrated on either side of the dorsal median line and at the distal margin of nota and tergites, with larger patches occurring laterally. Pat- terning mirrored on either side of the dorsal median line and continuing in this manner down the body; in some specimens, mid brown colouration not widespread past thoracic nota, leaving the abdominal tergites largely beige or ochre. Legs Legs generally ochreous and interspersed with beige pat- terning, the hind femora bearing striations laterally. Hind tibia ochreous with equidistant, pale beige spots dorsally, typically coinciding with pairs of larger dorsal linear spines. Fore and mid femur without ventral linear spines. Ventral surface of fore and mid tibia armed with two rows, bearing an average of three linear spines each, one prolateral and one retrolateral. Hind femur bearing an average of 14 (min. 8, max. 20) prolateral and 9 (6– 13) retrolateral ventral linear spines on inferior surface in males and 5 (0–13) and 8 (3–13) in females. Hind tibia with an average of 46 (35–57) prolateral and 37 (29–53) retrolateral dorsal linear spines in both sexes. First segment of hind tarsus with an average of 4 (2–6) dorsal linear spines prolaterally and 2 (0–4) ret- rolaterally. Second tarsal segment bearing an average of 2 dorsal linear spines, 1 (1–2) each prolaterally and retrolaterally. Male terminalia Suranal plate predominantly light to mid brown; glabrous except the distal margin, which is sparsely clothed with short setae; distal margin of plate strongly emarginate. Subgenital plate light brown and sparsely clothed with 10 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense https://www.inaturalist.org/observations/184652560 short brown setae; a rounded, poorly pronounced median lobe at the distal margin, longer than lateral lobes. Female terminalia Suranal plate beige medianly and mid brown laterally. Proximal margin of plate rounded, distal margin emargin- ate with a median notch. Entire surface of plate sparsely covered in short, brown setae and with longer setae densely concentrated at the distal margin. Subgenital sclerotised at the distal margin and bearing three beige, subtriangular lobes facing distally; median lobe shorter than lateral lobes. Ovipositor bearing eight moderately produced teeth. Remarks Speleotettix binoomea is the most morphologically and genetically divergent described species in the genus. It also exhibits variation in body colouration between populations, with males more prominently ochreous at Jenolan compared with Wombeyan (Figure 6). Beasley- Hall et al. (2025) also recovered crickets at Abercrombie as members of S. binoomea, but no material was available to us from that location for morphological examination. Etymology From binoomea (pronounced BIH-noo-mee), a Gundungurra word meaning “dark places” used to refer to the caves at Jenolan. The Aboriginal people of the Gundungurra Nation are the Traditional Custodians of the land on which the Jenolan Karst Conservation Reserve stands. The species name is an indeclinable noun in apposition. Distribution In caves in central and south-east NSW, Australia. Presently known from limestone and marble caves at Jenolan, Abercrombie and Wombeyan Karst Conservation Reserves. Speleotettix chopardi (Karny, 1935) nomen dubium (Figure 7) Pachyrhamma chopardi Karny, 1935, pp. 378–383, fig. 43. Speleotettix chopardi Chopard, 1944, p. 52 Material examined Holotype ?Juvenile ♂; Dandenong Ranges, VIC; 1935; coll. H. H. Karny 1935; MNHN EO-ENSIF4922. Examined as a digitised specimen via https://science.mnhn.fr/taxon/ species/pachyrhamma/chopardi. Remarks Karny described P. chopardi from a single male individual he suspected was a juvenile (Karny 1935). Karny did not provide a precise type locality for the species and did not document exact values for most of the morphological characters he examined. Chopard transferred the species to Speleotettix with the establishment of the genus, but it is still considered a member of Pachyrhamma in several checklists due to the original description being in French (ABRS 2023; Mesa 1970; Richards 1967). We have inspected the digitised holotype of S. chopardi and can confirm Karny’s measurements of the length of the body and leg segments are correct, and his approximate counts of the linear spines on the legs agree F I G U R E 6 Speleotettix binoomea Beasley-Hall, sp. nov. in life. The ochreous body colouration is prominent in males at Jenolan and evident to a lesser extent at Wombeyan (see male beneath the female in the mating pair). Images © Anne Musser and Helen Smith. REVISION OF THE CAVE CRICKET Speleotettix 11 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense https://science.mnhn.fr/taxon/species/pachyrhamma/chopardi https://science.mnhn.fr/taxon/species/pachyrhamma/chopardi with ours. The specimen appears to be a member of Spe- leotettix due to the lack of anterolateral spine on the fore coxae and the shape of the male suranal and subgenital plates. However, the holotype has distorted over time: the male suranal plate is very different to the original illustration by Karny, which depicts a weaker, U-shaped emargination at the distal margin more suggestive of Cavernotettix (Karny, 1935). Further, if the specimen is actually a juvenile, these characters may not be helpful in providing a generic identification in the first place. Based on our observations, Speleotettix adults are usually 11 mm or longer and Karny’s specimen is on the cusp at 11.5 mm. Karny states that the fore coxa of the specimen is unarmed (which would rule out Cavernotettix), but as he only examined a single individual, we cannot rule out the character being broken or simply missed. The female is not known. Karny’s lack of information regarding the specimen’s collection locality only complicates matters further. The Dandenong Ranges is a 35-km2 region of thick rainforest with elevations of up to 600 m and is a cool, mesic habi- tat typical of Australian Rhaphidophoridae. An unde- scribed species of Cavernotettix is found in the Ranges and a new genus, Eburnocauda Beasley-Hall and Iannello, 2024, was recently described only 30 km away (Iannello & Beasley-Hall 2024). Mesa (1970) and Richards (1987) referred to further undescribed species richness in the area, but this material was never deposited in a collection for examination. Given the high biodiversity of the region, we cannot rule out the existence of additional, undescribed members of Macropathinae being sympatric with S. chopardi. In other words, further sampling effort targeting the Dandenong Ranges may not resolve this problem. Considering the myriad of issues above, is our view that the quality of the description (and holotype) of S. chopardi currently makes its confident identification to the genus or species level an impossibility. In order to avoid future impediment to taxonomic work on the group in Australia, we believe S. chopardi represents a nomen dubium and a neotype will need to be designated in future. Distribution Dandenong Ranges, VIC, Australia. Precise locality unknown. F I G U R E 7 Speleotettix chopardi (Karny, 1935), holotype male. Specimen label (a); head (b); dorsal and ventral views of male terminalia (c,d); dorsal view of hind tibia (e). Images reproduced under a CC BY-NC-ND 4.0 licence (https://science.mnhn.fr/taxon/species/pachyrhamma/chopardi). Scale bar = 1 mm. Images without scale bars lacked them in the original photograph. 12 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense https://science.mnhn.fr/taxon/species/pachyrhamma/chopardi Speleotettix craggiensis (Richards, 1974) comb. nov. (Figure 8) Cavernotettix craggiensis Richards, 1974, pp. 256–259. Material examined Holotype Adult ♂; under boulders on Craggy Island, Bass Strait, TAS; coll. M. H. Christie, 18 June 1972; ANIC 14-042006. Paratypes Adult ♀; under stones on Craggy Island, Bass Strait, TAS; coll. J. S. Whinray, 16 June 1972; TMAG F000260. 1 ♀; same collection and locality information as previous; 19 June 1972; TMAG F000261. Sex not recorded; under stones on Craggy Island, Bass Strait, TAS; coll. M. H. Christie, ANIC 14-008962 (molecular data only). Diagnosis Speleotettix craggiensis can be differentiated from other members of Speleotettix by the female terminalia. The subgenital plate has distinctive square lateral lobes that are abruptly truncated at the distal margin, but not dark- ened by sclerotisation relative to the remainder of the plate. The suranal plate has a straight distal margin and the ovipositor has eight moderately produced teeth on the dorsal margin. Speleotettix craggiensis is the largest recorded member of Speleotettix, with a body size of up to 19 mm in males and 18 mm in females. Redescription As in Richards (1974), with exceptions as follows: hind femur bearing an average of 43 (min. 31, max. 59) ventral prolat- eral and 62 (48–79) retrolateral linear spines on inferior sur- face in males and 24 (18–35) and 35 (25–48) in females. Hind tibia with an average of 40 (32–47) prolateral and 42 (34–51) retrolateral dorsal linear spines in both sexes. The first segment of hind tarsus with an average of 3 (1–4) dorsal linear spines prolaterally and 4 (2–5) retrolaterally. F I G U R E 8 Speleotettix craggiensis (Richards, 1974) comb. nov. holotype and paratype. Head (a); dorsal view of terminalia (b ♂, d ♀); ventral view of male and female terminalia (c ♂ with subgenital plate removed, e ♀); lateral view of ovipositor (f,g); dorsal view of hind tibia (h). Scale bar = 1 mm. Male holotype (b,c) artificially paler than in life due to the age and preservation of the specimen. REVISION OF THE CAVE CRICKET Speleotettix 13 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense Second tarsal segment bearing an average of 2 dorsal linear spines, 1 (1–3) each prolaterally and retrolaterally. Remarks Richards (1974) incorrectly documented linear spine counts for this species in her original description. Macropathines typically bear a greater number of dorsal linear spines on the prolateral surface of the hind leg com- pared with the retrolateral, but counts in Richards’ original description are reversed in this respect. Upon inspecting type material of the species, we confirm this mistake and provide corrections to her original counts above. Values for the left and right side of the body have been averaged and rounded to the nearest whole number. Speleotettix craggiensis comb. nov. was described within Cavernotettix based on presumed affinities with Cavernotettix wyanbenensis Richards, 1966 and Cavernotettix buchanensis Richards, 1966. However, the species lacks the anterolateral spine on the fore coxae, a trait present in all other Cavernotettix and absent in Speleotettix. Additional characters shared by Speleotettix species include a greater number of teeth on the ventral valve of the ovipositor compared with Cavernotettix and prominent lateral lobes on the male suranal plate sepa- rated by a deep emargination at the distal margin. The spe- cies was recovered within Speleotettix in a recent molecular phylogeny by Beasley-Hall et al. (2025) (Figure 1), confirm- ing its generic placement. Speleotettix craggiensis comb. nov. is the only member of the genus currently known from TAS. The species is listed as rare under state environ- mental legislation owing to its extremely limited distribu- tion (Threatened Species Section 2022). Distribution Craggy Island, Bass Strait, TAS, Australia. Recorded under boulders and in burrows of breeding seabirds (Richards 1974, 1987). Speleotettix palaga Beasley-Hall, sp. nov. (Figure 9) http://zoobank.org/urn:lsid:act:34BE5EB6-9E3E-496D- 8169-78372DA09AF0 Speleotettix sp. nov. 2 (Beasley-Hall et al., 2025). Material examined Holotype Adult ♂, unnamed mining tunnel in Wombat-Lerderderg National Park near Blackwood, VIC, �37.480278, 144.330222, coll. S. Iannello, 16 December 2023, ANIC 14-009162. Paratypes 2 ♀, 2 ♂ (all subadult), same locality and collection infor- mation as holotype, ANIC 14-009163–14-009166. Other material 2♀, 1 ♂, Imperial Mine, Jack Cann Reserve in Wombat State Forest, VIC, �37.478694, 144.290833, coll. S. Iannello, 16 December 2023, ANIC 14-009167–14-009169. 1 ♀, unnamed mining tunnel in Blackwood, VIC (photograph only), �37.478444, 144.290972, iNaturalist 27853615. 1 ♀, Bullarto, VIC (photograph only), �37.404722, 144.218111, iNaturalist 213067284. 1 ♀, Lerderderg area, VIC (photograph only), �37.492278, 144.436389, iNaturalist 125136101. 1 ♂, Blighs Road near Lyonville, VIC (photograph only), �37.386056, 144.288472, iNaturalist 173174834. 1 ♂, Domino Road near Lyonville, VIC (photograph only), �37.397667, 144.278028, iNaturalist 204638088. Diagnosis Speleotettix palaga can be differentiated from other mem- bers of Speleotettix by its body colouration and the female terminalia. Adults often have a row of prominent tuber- cles on the distal margin of each of the thoracic nota and abdominal tergites (Figure 10, bottom image). The lateral lobes of the subgenital plate are subrectangular with rounded corners and a mid brown, distinct from the remainder of the plate. The ovipositor has six moderately produced teeth on the dorsal margin. Description Measurements Holotype body length 16 mm, hind tibia 22 mm. ♀ para- type 13 mm, hind tibia 18 mm, ovipositor 10 mm. Head Mid brown banding on vertex and frons of head forming a bold M shape, extending to a horizontal band spanning fastigium. Body Body varying from predominantly mid brown to beige, with brown mottling covering thoracic nota and abdominal tergites and interrupted by beige patches. Small, well-defined spots generally concentrated on either side of the dorsal median line and at the distal margin of nota and tergites, with larger patches occurring laterally. Patterning mirrored on either side of the dorsal median line and continuing in this manner down the body. Abdomen often with a row of prominent beige tubercles at distal margins of tergites, appearing similar to raindrops clinging to the body. 14 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense http://zoobank.org/urn:lsid:act:34BE5EB6-9E3E-496D-8169-78372DA09AF0 http://zoobank.org/urn:lsid:act:34BE5EB6-9E3E-496D-8169-78372DA09AF0 https://www.inaturalist.org/observations/27853615 https://www.inaturalist.org/observations/213067284 https://www.inaturalist.org/observations/125136101 https://www.inaturalist.org/observations/173174834 https://www.inaturalist.org/observations/204638088 Legs Legs generally light brown and interspersed with beige patches, with hind femora also bearing beige striations laterally. Hind tibia with equidistant, pale beige spots dor- sally, typically coinciding with pairs of larger dorsal linear spines. Fore and mid femur without ventral linear spines. Ven- tral surface of fore and mid tibia armed with two rows, bearing an average of three linear spines each, one pro- lateral and one retrolateral. Ventral surface of hind femur bearing an average of 32 (min. 18, max. 60) prolateral and 29 (11–49) retrolateral linear spines in males and 16 (0– 38) and 18 (8–34) in females. Hind tibia with an average of 40 prolateral and 37 retrolateral dorsal linear spines in both sexes. First segment of hind tarsus with an average of 3 (2–4) dorsal linear spines prolaterally and 3 (2–5) retrolaterally. Second tarsal segment bearing an average of 2 dorsal linear spines, 1 (1–2) prolaterally and 1 (no variation) retrolaterally. Male terminalia Suranal plate predominantly mid brown and glabrous except the distal margin, which is sparsely clothed with short setae; distal margin of plate strongly emarginate. Subgenital plate light brown and sparsely clothed with short brown setae; a rounded, poorly pronounced median lobe at the distal margin, longer than lateral lobes. Female terminalia Suranal plate ranging from uniformly dark brown to dark brown patterning restricted to lateral regions only. Proxi- mal margin of plate rounded, distal margin emarginate. Entire surface of plate sparsely covered in short, brown setae and with longer setae densely concentrated at the distal margin. Subgenital plate sclerotised at the distal margin; thickening of cuticle particularly pronounced at the lateral lobes, which are mid to dark brown and sub- rectangular; median lobe very well pronounced with a tri- angular apex and longer than lateral lobes. Ovipositor bearing six moderately produced teeth. Etymology From the Latin palaga, meaning gold ingot, in reference to the disused gold mines of the Blackwood area in which the species is found. It is a feminine noun used in apposition. F I G U R E 9 Speleotettix palaga Beasley-Hall, sp. nov. head (a); dorsal view of terminalia (b ♂, d ♀); ventral view of male and female terminalia (c ♂, e ♀); lateral view of ovipositor (f,g); dorsal view of hind tibia and tarsus (h). Scale bar = 1 mm. REVISION OF THE CAVE CRICKET Speleotettix 15 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense Distribution Known from mining tunnels in the Blackwood area near and within Wombat–Lerderderg State Park, VIC, Australia. Speleotettix tindalei Chopard, 1944 Speleotettix tindalei Chopard, 1944, pp. 53–54. (Figure 11) Material examined Cotypes 1 ♂, 1 ♀; cave section 272, Hindmarsh S. E. [Tantanoola Caves], SA; coll. N. B. Tindale [n.d.]; MNHN EO-ENSIF4918 and EO-ENSIF4919. Examined as digitised specimens via https://science.mnhn.fr/taxon/species/speleotettix/tindalei. Other material 3 ♀, 1 ♂; toilet block at Tantanoola Caves Conservation Park, SA; collection date unknown, provided to the authors on 19 March 2023, ANIC 14-009172–14-009175. 1 ♀, 3 ♂; on road at Donovans, SA near VIC border; �38.015194, 140.958278; coll. S. Bourne, 1 May 2016. 1 ♀; ‘Fox Hole’ Cave near Monbulla, SA; �37.419556, 140.677111; coll. P. Beasley-Hall and A. D. Austin, 21 May 2016. 4 ♀, 2 ♂; Tantanoola Caves Conservation Park; iNa- turalist 202122890, 184652288, 148238575, 127656272, 112495949, 133543241 (photographs). 1 ♂; unnamed cave on private property near Mount Schank; �37.924138, 140.676520; iNaturalist 221982389 (photo- graph). ♂; Moerlong area, approx. 9 km SSE from ‘Fox Hole’ Cave; �37.490974, 140.736310; iNaturalist 185713623 (photograph). Diagnosis The female subgenital plate of S. tindalei resembles that of S. palaga in that the lateral lobes are strongly sclerotised and darkened in both species, but in tindalei the lobes are not pronounced and are darkened at the distal margin only. In S. palaga, the dark brown colouration extends to the median surface of the lobes, which are rounded and jut out from the surrounding plate. The ovipositor has eight weakly produced teeth on the dorsal margin. While the trait exhibits considerable variation within species, it is also noteworthy that S. tindalei has comparatively modest counts of ventral linear spines of the hind femur: we have not observed more than 36 spines on the prolateral or ret- rolateral surfaces. In other members of Speleotettix, these counts may reach 60 (e.g., in S. aolae and palaga). These counts are lower only in S. binoomea (20 or fewer). Redescription Measurements Body length up to 16 mm, hind tibia up to 29.5 mm; ovi- positor up to 13 mm. Head Mid brown banding on vertex and frons of head forming a rough M shape, extending to a horizontal band span- ning fastigium. Body Body generally mid brown and beige, with brown mot- tling covering thoracic nota and abdominal tergites and interrupted by beige patches. Small, well-defined spots generally concentrated on either side of the dorsal median line and occasionally at the distal margin of nota and tergites, with larger patches occurring laterally. Pat- terning mirrored on either side of the dorsal median line and continuing in this manner down the body. Legs Legs generally light brown and interspersed with beige patches, with the hind femora also bearing beige F I G U R E 1 0 Speleotettix palaga Beasley-Hall, in life, with distinct raindrop-like tubercles at the distal margins of tergites (bottom image). Images © Paul George and user Mononymous via iNaturalist (observations 27853615, 204638088), reproduced under Creative Commons licences. 16 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense https://science.mnhn.fr/taxon/species/speleotettix/tindalei https://www.inaturalist.org/observations/202122890 https://www.inaturalist.org/observations/184652288 https://www.inaturalist.org/observations/148238575 https://www.inaturalist.org/observations/127656272 https://www.inaturalist.org/observations/112495949 https://www.inaturalist.org/observations/133543241 https://www.inaturalist.org/observations/221982389 https://www.inaturalist.org/observations/185713623 striations laterally. Hind tibia with equidistant, pale beige spots dorsally, typically coinciding with pairs of larger dor- sal linear spines. Ventral surface of fore and mid tibia armed with two rows, bearing an average of three linear spines each, one prolateral and one retrolateral. Ventral surface of hind femur bearing an average of 21 (10–36) prolateral and 17 (9–28) retrolateral linear spines in males and 27 (19– 36) and 23 (17–28) in females. Hind tibia with an average of 52 (45–59) prolateral and 48 (41–57) retrolateral dorsal linear spines in both sexes. First segment of hind tarsus with an average of 4 (2–6) dorsal linear spines prolaterally and 3 (2–4) retrolaterally. Second tarsal segment bearing an average of 2 dorsal linear spines, 1 (0–2) prolaterally and 1 (no variation) retrolaterally, averages in agreement with Chopard’s counts. Male terminalia Suranal plate predominantly light brown; glabrous except the distal margin, which is sparsely clothed with short setae; distal margin of plate strongly emarginate. Subge- nital plate light brown and sparsely clothed with short brown setae; a rounded, poorly pronounced median lobe at the distal margin, longer than lateral lobes. Female terminalia Suranal plate beige medianly and mid to dark brown lat- erally. Proximal margin of plate rounded, distal margin weakly emarginate. Entire surface of plate very sparsely covered in short, pale setae and with longer setae densely concentrated distolaterally. Subgenital plate sclerotised with lateral lobes truncate, thick and darkened at the dis- tal margin; median lobe well pronounced, longer than lat- eral lobes and ending at a pointed apex; together, the three lobes resemble a curly bracket. Remarks Speleotettix tindalei was described by Chopard in 1944 and is the type species of the genus. While it is clearly nested within Speleotettix based on molecular (Beasley- Hall et al. 2025) and morphological data presented here, Chopard did not provide precise numbers of spines on F I G U R E 1 1 Speleotettix tindalei Chopard, 1944. Head (a); dorsal view of terminalia (b ♂, d ♀); ventral view of male and female terminalia (c ♂, e ♀); lateral view of ovipositor (f,g); distal view of hind tibia (h). Scale bar = 1 mm. REVISION OF THE CAVE CRICKET Speleotettix 17 of 20 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense segments of the hind legs to facilitate generic identifica- tion and instead detailed approximate values. Chopard’s description of the terminalia is also vague and limited to the male suranal and female subgenital plates. Counts of spines from fresh specimens collected from the species’ type locality broadly agree with those pro- vided by Chopard (e.g., the dorsal surface of the hind tibia bearing 41–59 linear spines on either side compared with Chopard’s ‘around 50’). However, we count eight teeth on the ventral surface of the ovipositor after examining fresh specimens and digitised type material, whereas Chopard recorded six. This is likely due to the most proxi- mal and distal teeth not being preceded by a ridge on the median surface, making them difficult to see. We note that Chopard named the type locality of the species as “Hindmarsh S.E.” referring to the Hundred of Hindmarsh, an administrative division overlapping with Tantanoola Caves Conservation Park. Hindmarsh is also a suburb in metropolitan Adelaide, almost 400 km north- west of this locality. This seems to have confused Richards, who alluded to a disjunct but unnamed popula- tion of Speleotettix (presumably S. tindalei) in the Adelaide region in addition to this Limestone Coast population. To our knowledge, no Rhaphidophoridae have been docu- mented between the Limestone Coast and Nullarbor Plain in SA, and we consider this an error on Richards’ part. Distribution Caves in the Limestone Coast of SA (Tantanoola, Naracoorte, and near Monbulla) and, according to Mesa (1970), across the nearby Victorian border at Princess Margaret Rose Cave. DISCUSSION The increase in the number of Speleotettix species described here represents a significant expansion of the geographic range of the genus, previously only known from two discrete locations �450 km apart in the south- east of SA and near the Melbourne metropolitan area of VIC (Figure 1). The new species can be delimited using a combination of characters relevant to body colouration, the suranal and subgenital plates, and the ovipositor. However, because considerable intraspecific variability is observed in species of Macropathinae (Richards 1958, 1971), we chose to supplement our descriptions with molecular evidence generated here and in a recent phy- logeny (Beasley-Hall et al. 2025) (Figure 1). In our previous phylogenetic study, we sampled a subset of the speci- mens examined here (with the exception of S. chopardi, see Taxonomy section) and recovered Speleotettix as reciprocally monophyletic with its sister genus, Caverno- tettix (Figure 1). Cavernotettix craggiensis from the Bass Strait of TAS was also recovered within Speleotettix with high node support, in agreement with its morphological similarity with other members of the genus. Our species descriptions are also supported by several distance and tree-based delimitation methods summarised in Figure 3. All species for which COI data were available were sepa- rated by at least 4.4% pairwise genetic divergence (Figure 3, Table S1), in agreement with a putative barcode gap in other Rhaphidophoridae (>2.5%) (Allegrucci et al. 2021; Zhao et al. 2022) and the Orthoptera more broadly (>3%) (Huang et al. 2013; Timm et al. 2022; Zhou et al. 2019). An additional finding from the phylogeny by Beasley- Hall et al. (2025) is the suggestion of an additional unde- scribed species found at Wee Jasper Caves in NSW. This lineage was recovered as sister to S. binoomea in that study, a split supported here by formal delimitation ana- lyses (Figures 1 and 3). However, we do not have adult material of this species available for examination to facili- tate a description. Specimens from Wee Jasper were deposited by Richards as additional “types” of Speleotettix profundus alongside the Jenolan, Wombeyan, and Abercrombie populations examined here (Figures 1 and 6). Richards also deposited material from caves in Bungonia approximately 50 km south of Wombeyan under the same tag name, which may represent an additional population of S. binoomea. Ultimately, in the present study we have resolved the genus and species-level status of Speleotettix profundus, which until now had remained enigmatic for over 60 years. Despite the increased known distribution of the genus, we note most species of Speleotettix can be classi- fied as short-range endemics under criteria set out by Harvey (2002), defined as naturally inhabiting a range of <10 000 km2. Short-range endemics are particularly vul- nerable to extinction and may require conservation listing to ensure their protection in the long term. At present the only protected species in the genus is S. craggiensis, known exclusively from under boulders and the burrows of breeding seabirds on Craggy Island, a small rocky island about 20 km from the tip of Flinders Island in the Bass Strait (Richards 1974). The species is listed as rare under the Tasmanian Threatened Species Protection Act 1995 and has no conservation management plan in place. Threats to all Speleotettix species, not just craggiensis, include clearing of native forest, predation by invasives, and human disturbance via tourism. Conservation listing is also necessary for these remaining species because they may have narrower habitat or temperature toler- ances compared with epigean relatives, and therefore a greater vulnerability to anthropogenic disturbance and climate change (Mammola et al. 2019). ACKNOWLEDGEMENTS We would like to give a hearty thank you to Sil Iannello (Australian Speleological Federation), Helen Smith (AM) and Anne Musser and Michael Collins (Jenolan Caves Reserve Trust) for collecting and transporting specimens. 18 of 20 BEASLEY-HALL ET AL. 20521758, 2025, 3, D ow nloaded from https://onlinelibrary.w iley.com /doi/10.1111/aen.70011 by M assey U niversity L ibrary, W iley O nline L ibrary on [29/07/2025]. See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense Thank you also to Andreas Zwick and You Ning Su (ANIC), Derek Smith (AM), and Karin Koch (QM) for facilitating loans of museum material. We are grateful to Gundun- gurra Elder Auntie Sharyn Halls, who approved the use of the S. binoomea species name, and Andrew Le Lievre (Jenolan Caves Reserve Trust) for liaising with Auntie Sharyn. Finally, a big cheers to Mark Harvey (Western Aus- tralian Museum) for providing helpful advice regarding the International Code on Zoological Nomenclature. Open access publishing facilitated by The University of Adelaide, as part of the Wiley - The University of Adelaide agree- ment via the Council of Australian University Librarians. CONFLICT OF INTEREST STATEMENT The authors have no conflict of interest to disclose. DATA AVAILABILITY STATEMENT Newly generated COI data from the Wombeyan popula- tion of Speleotettix binoomea are available via the Gen- Bank repository and accessions are listed under Material Examined for that species. Specimens were collected under the following permits: AA-0001076 via Parks Victoria and FS/14-3694/1_2024 via the Victorian Department of Energy, Environment and Climate Action, both held by PGBH (Victoria); Y27154-2 via the South Australian Department for Environment and Water, held by ER (South Australia); and SL102683 via the New South Wales Department of Planning and Environment, held by the Australian Museum (New South Wales). ORCID Perry G. Beasley-Hall https://orcid.org/0000-0002-7360- 7933 REFERENCES ABRS. (2023). Species Pachyrhamma chopardi Karny, 1935. Australian Faunal Directory. Available from: http://biodiversity.org.au/afd. publication/8524b8f3-eedb-464f-a97b-6bd91a943983 [Accessed 21 June 2023]. 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See the T erm s and C onditions (https://onlinelibrary.w iley.com /term s-and-conditions) on W iley O nline L ibrary for rules of use; O A articles are governed by the applicable C reative C om m ons L icense https://doi.org/10.1093/biosci/biz064 https://doi.org/10.1111/1755-0998.13281 https://doi.org/10.11646/zootaxa.4154.3.9 https://doi.org/10.11646/zootaxa.4154.3.9 https://doi.org/10.1111/j.1558-5646.2007.00023.x https://doi.org/10.1139/cjz-2022-0041 https://doi.org/10.1080/03014223.1999.9517590 https://doi.org/10.1080/03014223.1999.9517590 https://doi.org/10.1016/j.jnc.2017.01.003 https://doi.org/10.1016/j.jnc.2017.01.003 https://doi.org/10.1093/bioinformatics/btt499 https://doi.org/10.1093/bioinformatics/btt499 https://doi.org/10.3897/zookeys.1123.86704 https://doi.org/10.3897/zookeys.1123.86704 https://doi.org/10.3897/zookeys.1109.73937 https://doi.org/10.3897/zookeys.1109.73937 https://doi.org/10.1186/s12862-019-1404-5 https://doi.org/10.1111/aen.70011 https://doi.org/10.1111/aen.70011 Integrative taxonomic revision of the Australian cave cricket Speleotettix Chopard, 1944 (Orthoptera: Rhaphidophoridae): Ne... Abstract INTRODUCTION MATERIALS AND METHODS Taxon sampling and imaging DNA sequencing and phylogenetic analysis Species delimitation Abbreviations and terminology Collection abbreviations Locations Morphological terminology RESULTS Species delimitation TAXONOMY Genus Speleotettix Chopard, 1944. Diagnosis Redescription Head Body Legs Male terminalia Female terminalia Remarks Etymology Distribution Key to species of Speleotettix Speleotettix aolae Beasley‐Hall, sp. nov. Material examined Holotype Paratype Other material Diagnosis Description Measurements Head Body Legs Male terminalia Female terminalia Remarks Etymology Distribution Speleotettix binoomea Beasley‐Hall, sp. nov. Material examined Holotype Paratypes Diagnosis Description Measurements Head Body Legs Male terminalia Female terminalia Remarks Etymology Distribution Speleotettix chopardi (Karny, 1935) nomen dubium Material examined Holotype Remarks Distribution Speleotettix craggiensis (Richards, 1974) comb. nov. Material examined Holotype Paratypes Diagnosis Redescription Remarks Distribution Speleotettix palaga Beasley‐Hall, sp. nov. Material examined Holotype Paratypes Other material Diagnosis Description Measurements Head Body Legs Male terminalia Female terminalia Etymology Distribution Speleotettix tindalei Chopard, 1944 Material examined Cotypes Other material Diagnosis Redescription Measurements Head Body Legs Male terminalia Female terminalia Remarks Distribution DISCUSSION ACKNOWLEDGEMENTS CONFLICT OF INTEREST STATEMENT DATA AVAILABILITY STATEMENT ORCID REFERENCES SUPPORTING INFORMATION