First records of Hippasosa ghost (Jocque & Jocqué, 2017) (Araneae, Lycosidae) outside of the type locality in northwestern Madagascar

the journal of biodiversity data NOTES ON GEOGRAPHIC DISTRIBUTION 1257 Academic editor: Thiago Moreira Received: 23 October 2025 Accepted: 29 November 2025 Published: 17 December 2025 Copyright © The authors. This is an open‑access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0) Abstract. The Malagasy endemic wolf spider Hippasosa ghost (Jocque & Jocqué, 2017), previously known from a single small patch of sandy habitat adjacent to a permanent lake in the Mariarano Classified Forest in northwestern Madagascar, is recorded from two additional localities with similar habitat along the Mariarano River, 10 km and 45 km from the type locality. These finds indicate the possibility that this highly specialized species could have a larger distribution than originally thought, but also they show the importance of protecting such vulnerable habitats. Key words. Dry tropical forest, Mariarano, psammophilous, specialist Pett BL, Escobar-Toledo J, de Beer L, Murray KI, Rakotorahalahy M, Siméon F, Rivoson JL (2025) First records of Hippasosa ghost (Jocque & Jocqué, 2017) (Araneae, Lycosidae) outside of the type locality in northwestern Madagascar. Check List 21 (6): 1257–1261. https://doi.org/10.15560/21.6.1257 Introduction The wolf spider (Araneae, Lycosidae) genus Hippasosa Roewer, 1960 has a relatively wide distribution, with 13 species distributed across Africa (including Madagascar), Arabia, and China. Revised under their previous generic name of Ocyale Audouin, 1826 by Alderweireldt (1996), the genus was declared a nomen dubium by Sherwood (2022), and all species were transferred to Hippasosa. Hippasosa are diagnosed by two elongated curved prongs on the palea, epigyne with white hairs and a wide inverted T-shaped sclerite, large globular spermathecae and the conformation of the copulatory ducts (Alderweireldt 1996; Jocque et al. 2017). Hippasosa ghost was described in Ocyale by Jocque & Jocqué (2017) as the first spider discovery from a biomonitoring program in the dry tropical forest ecoregion of Mariarano classified forest, Mahajanga (NW Madagascar) (Figure 1A), from which several others have been described in recent years (e.g. Pett and Rabemananjara 2022; Escobar-Toledo and Pett 2024; Murray et al. 2024). The species is only known from a single white-sand beach (< 2 ha) in the middle of dry scrub forest adjacent to a permanent lake. Individuals are remarkable in their psammophilous habits, creating retreats and burrows lined with silk to shelter from the heat of the day. Burrows are 30–50 cm deep and 15–20 °C cooler than the sand surface (BLP unpublished data). Pett and Bailey (2019) studied the ecology of H. ghost at the type locality using a patch occupancy modelling framework; finding that occupancy probability was much higher closer to the adjacent lake, on bare sand rather than areas with low vegetation or trees, and some positive association with west-facing slopes. The population is also likely close to just under ~1,000 adult individuals at any one time. Pett had visited 14 candidate sandy patch sites in the general region (within 30 km) and found no individuals of H. ghost at any site. Considering the conspicuousness of the species, we believe those to be true absences likely related to the properties of the sand. Pett and Bailey (2019) commented on the possible conservation status of, and threats to, this enigmatic spider and considered it to be a possible 21 (6) · https://doi.org/10.15560/21.6.1257 21 (6): 1257–1261. https://doi.org/10.15560/21.6.1257 First records of Hippasosa ghost (Jocque & Jocqué, 2017) (Araneae, Lycosidae) outside of the type locality in northwestern Madagascar Brogan L. Pett1 , 2 , 3 , Jaime Escobar-Toledo1, 3 , Lenon de Beer4, Katie I. Murray1, 3 , 5 , Mamitiana Rakotorahalahy6, FandresenaSiméon6, Jacquis L. Rivoson7 1 SpiDiverse, Biodiversity Inventory for Conservation NPO (BINCO), 3380 Walmersumstraat, Glabbeek, Belgium 2 University of Iceland, Faculty of Life and Environmental Sciences, Sturlugata 7, 102 Reykjavik, Iceland 3 Operation Wallacea, Lincolnshire, UK 4 Eden People plus Planet, Antananarivo Madagascar 5 Centre for Ecology and Conservation, College of Life and Environmental Sciences, University of Exeter, Penryn Campus, Penryn, Cornwall, TR10 9FE, UK 6 University of Antananarivo, Faculty of Sciences, Department of Entomology. Mention ECES (Entomologie Culture Élevage et Santé) 7 Faculty of Science, Technology and the Environment (FSTE), University of Mahajanga, Madagascar Corresponding authors: Brogan L. Pett ([email protected]); Katie I. Murray ([email protected]) Check List 21 (6) · https://doi.org/10.15560/21.6.1257 Pett et al. · New records of Hippasosa ghost 1258 single-site endemic with considerable vulnerability to stochastic events or climatic shift. Several years later, further samples from the same general forest site but separate camp yielded several individuals of Hippasosa, and additionally a population of Hippasosa was discovered by LdB in the region. Taxonomic bias in research (Clark and May 2002) means that the Wallacean shortfall in invertebrate conservation (discussed comprehensively by Cardoso et al. 2011) is as pressing issue as any and is not improving. Thus, it is imperative that biodiversity records of little-studied species be published in the scientific literature to contribute towards understanding of species’ ecology and distribution. As a pertinent Malagasy example, only 47 of the 108 salticid species recorded from Madagascar have been reported in the past 100 years (Murray et al. 2024). The aims of this study are therefore to report on the identity of the aforementioned Hippasosa specimens as H. ghost and to illustrate this species, present a new distribution map, and briefly discuss its conservation and distribution status. Materials and methods Spiders were collected during expeditions to the Mariarano classified forest. These expeditions were co-ordinated by Biodiversity Inventory for Conservation NPO (BINCO) in partnership with Operation Wallacea. All spiders were collected by dedicated active nocturnal searching with headlamps. All material is preserved in 70% ethanol. The left pedipalp of a male was dissected and illustrated. Examinations were carried out with an AmScope ZM-4T stereomicroscope or an Olympus BX61. Images were taken using a Visionary Digital BK Laboratory System using a Canon 5D camera and a 65-mm macro zoom lens. All images are composites of z-slices of 20–30 images, compiled using the program Helicon Focus v. 6.7. Images were adjusted in Adobe Photoshop v. 21.0.1 for contrast and white balance. Final figures were composed in Adobe Photoshop. All specimens have been deposited in the Royal Museum for Central Africa (RMCA), Tervuren, Figure 1. Map of known distribution and habitat of H. ghost. A. Map of records, including the type locality in the Matsedroy lakes (yellow circle) and the new records (blue triangles). B, C. Habitat at Papamena village. All specimens are collected from locations along the Mariarano River, except those at the more isolated type locality. Check List 21 (6) · https://doi.org/10.15560/21.6.1257 Pett et al. · New records of Hippasosa ghost 1259 Belgium. Export permit for biological samples was issued by the Ministère de l’Environnement et du Développement Duriable, permit no. 208N-EA08/MG24 for the Mahamavo–Mahajanga region. Results Araneae Clerck, 1757 Lycosidae Sundevall, 1833 Hippasosa Roewer, 1960 Hippasosa ghost (Jocque & Jocqué, 2017) Ocyale ghost Jocque & Jocqué, in Jocque et al. 2017: 5, figures 3–7 (holotype male, Mariarano classified forest, Mahajanga, Madagascar; RMCA.245337). Hippasosa ghost—Sherwood 2022: 583 (transfer from Ocyale). Figure 2 Figure 2. Hippasosa ghost male. A–D. Habitus. E. Male pedipalp ventral view. F. Male pedipalp prolateral view. A, B. In vivo. C. Specimen in dorsolateral view. D. specimen in dorsal view. Scale bars: D = 2 mm; E = 0.5 mm. Abbreviation: MA = median apophysis. Photo credit: LdB (A, B), BLP (C–F). Check List 21 (6) · https://doi.org/10.15560/21.6.1257 Pett et al. · New records of Hippasosa ghost 1260 New records. MADAGASCAR — Boeny region • Mahajanga, Mariarano, Mariarano river; −15.5062, 46.6977; elev. 90 m; 05–19.VII.2024; B. Pett, K. Murray, J. Escobar-Toledo, M. Rakotorahalahy & J. Rivison leg.; bare sandy riverbank; 3 ♂, 5 ♀, in ethanol, RMCA 247840 • Papamena village; −15.5758, 46.7386; 02.VI.2024; L. de Beer & B. Pett leg.; bare sandy riverbank; 1 ♂, 1 ♀, in ethanol, RMCA 247841. Diagnosis (from Jocque et al. 2017). Males are recognized by the distal prong of the palea appendage being much narrower than the proximal one and the median apophysis with the perpendicular prong is rectangular. Females are distinguished from other species of the genus by the epigyne, which is surrounded (not covered) by white setae, leaving the posterior sclerite exposed. Identification. Our specimens share the same highly conspicuous and regionally unique somatic morphology, being almost entirely white (females) or occasionally light brown (males) and with yellow eye rings. Genitalia match that of the type specimens, with the two-pronged palea with distal prong much narrower and identical posterolaterally curved MA for males (Figure 2E, F; compare with Jocque et al. 2017: figures 5B, 7C), and with the characteristic exposed epigyne for females. Distribution. Known only from the Mahajanga area (Boeny region, Madagascar), centred on Mariarano classified forest area. Now also known from two sites within 45 km of the type locality (Figure 2A). Habitat. Bare terrain with white-sand substrate on lakeshores and riverbanks (Figure 2B, C). Discussion Here, we report on the first two new localities for Hippsosa ghost outside of its type locality. One is only 10 km away, in the bank of the Mariarano River, whilst the other is around 45 km away in similar sandy environs. These records have important ramifications for our knowledge on the species, with H. ghost likely being a strong disperser as in some other psammophilous lycosids, e.g. Allocosa senex (Mello-Leitão, 1945)from South America, present in many sand-dune habitats in eastern and central South America (Postiglioni et al. 2019). The fact that this species has been found in similar habitats as the type locality along a nearby river, could indicate the importance of those natural corridors for the dispersion of the species. This could also mean that the species is perhaps much more widely distributed than even currently reported here and is not a single site endemic. Regardless, the species is clearly a habitat specialist and threatened by a unique set of circumstances (see Pett and Bailey 2019 for discussion). For example, the drought that gripped Madagascar in 2018–2021 considerably affected animals. The unique ecology of H. ghost, in creating silken retreats in burrowable sands near freshwater lakes likely makes it vulnerable to even slight changes in the moisture levels in sandy substrate (i.e. drying out). As specialists outcompete generalists in optimal habitat, and the inverse is true in disturbed or variable environments (Bonte et al. 2006), it is expected that climatic shift and droughts will favour the survival of generalist species as conditions become unstable. Hippasosa ghost outcompetes other lycosids and epigean spiders at the type locality, yet it is not present outside the immediate sandy patch around the lake (Pett and Bailey 2019). Therefore, whilst these new localities are important for the species conservation status, we urge restraint in dismissing H. ghost as of lower conservation relevance as its only habitat is a highly precarious one. Acknowledgements We would like to thank the people and guides living in and around the Mariarano community for their expert guidance and long-term collaboration with our project. Additionally, the students who joined the Operation Wallacea expeditions offered invaluable assistance to the project. Specimens were collected and exported under permit no. 208N-EA08/MG24. Additional information Conflict of interest The authors declare that no competing interests exist. Ethical statement No ethical statement is reported. Check List 21 (6) · https://doi.org/10.15560/21.6.1257 Pett et al. · New records of Hippasosa ghost 1261 Funding This study was financially supported by Operation Wallacea through the administration of the Mariarano field station. Author contributions Conceptualization: BLP, JET, LDB. Investigation: BLP, JET, LDB, KIM, MR, FS, JR. Methodology: BLP, JET, LDB, KIM, MR, FS, JR. Resources: BLP. Visualisation: BLP, LDB. Project administration: BLP. Writing – original draft: BLP, LDB. Writing – review and editing: JET, KIM, MR, JR. Map workLinda McKenzie (Digital Earth) Author ORCID iDs Brogan L. Pett https://orcid.org/0000-0002-0461-3715 Jaime Escobar-Toledo https://orcid.org/0009-0007-2495-780X Katie I. Murray https://orcid.org/0009-0007-2042-2285 Data availability All data that support the findings of this study are available in the main text. References Alderweireldt M(1996) A taxonomic revision of the genusOcyaleAudouin, 1826 in Africa (Araneae: Lycosidae).Journal of Natural History30 (9): 1349–1365. https://doi.org/10.1080/00222939600771261 Bonte D, Lens L, Maelfait J (2006) Sand dynamics in coastal dune landscapes constrain diversity and lifehistory characteristics of spiders. Journal of Applied Ecology 43 (4): 735–747. https://doi.org/10.1111/j.13652664.2006.01175.x Cardoso P, Erwin TL, Borges PAV, New TR (2011) The seven impediments in invertebrate conservation and how to overcome them. Biological Conservation 144 (11):2647–2655. https://doi.org/0.1016/j.biocon.2011.07.024 Clark JA, May RM (2002) Taxonomic bias in conservation research. Science 297 (5579): 191–192. https://doi.org/ 10.1126/science.297.5579.191b Escobar-Toledo J, Pett BL(2024) Contributions to theLarinia-group (Araneae: Araneidae) in Madagascar, with the description of two new species.Arachnology19 (9): 1312–1317. https://doi.org/10.13156/arac.2024.19.9.1312 Jocque M, Wellens S, Andrianarivosoa JD, Rakotondraparany F, The Seing S, Jocqué R (2017) A new species ofOcyale(Araneae, Lycosidae) from Madagascar, with first observations on the biology of a representative in the genus.European Journal of Taxonomy355. https://doi.org/10.5852/ejt.2017.355 Murray KI, Escobar-Toledo J, Pett, BL(2024) Three new species of plexippine jumping spiders (Salticidae, Salticinae, Plexippini) from dry forest in Boeny region, north-western Madagascar.African Invertebrates65 (2): 61–74. https://doi.org/10.3897/AfrInvertebr.65.126810 Pett BL, Bailey (2019) Ghost-busting: patch occupancy and habitat preferences of Ocyale ghost (Araneae: Lycosidae), a single site endemic in north-western Madagascar. Austral Entomology 58 (3): 1–11. https://doi.org/10.11 11/aen.12425 Pett BL, Rabemananjara PB(2022) A new species ofCopa(Araneae: Corinnidae: Castianeirinae) from dry forests in the north west of Madagascar.Zootaxa5115 (2): 281–287. https://doi.org/10.11646/zootaxa.5115.2.7 Postiglioni R, Bidegaray-Batista L, Simó M, Arnedo MA (2019) Move to stay: genetic structure and demographic history of a wolf spider inhabiting coastal sand dunes of southern South America. Systematics and Biodiversity 17 (7): 635–649. https://doi.org/10.1080/14772000.2019.1689197 Sherwood D (2022) On the taxonomic and nomenclatural status of Ocyale Audouin, 1826 (Araneae: Pisauridae) and Hippasosa Roewer, 1960 (Araneae: Lycosidae), with notes on some other taxa.Arachnology 19 (2): 582–584. https://doi.org/10.13156/arac.2022.19.2.582