Here, we present an overview of the global diversity of troglobionts. However, it is important to highlight that these numbers are subject to various biases: many are not highly accurate, do not adhere to rigorous criteria regarding the adopted concept of troglobiont, are significantly outdated, or rely on information that is unavailable or has not been compiled. Furthermore, several discrepancies among these estimates can be identified. We use them here as reference points for compilation, based on the most reliable data available.
We have highly consistent data for the former USSR (including Central Asia), Africa, Iran, Southeast Asia through New Guinea and Vanuatu, China, Canada and the USA, Mexico, Guatemala through Panama, the Caribbean (with Jamaica and Cuba treated separately), Venezuela, Colombia, Ecuador, Peru, Bolivia, and Brazil. We have less consistent data for Europe (particularly regarding Coleoptera and Diplopoda), the Middle East (excluding Iran), the Indian subcontinent, Japan, Australia, the smaller Pacific islands (excluding Vanuatu), New Zealand, the Guianas, and the Southern Cone of South America.
In October 23, 2026 our work believes that 9,559 troglobic documented in world (sum 9,563 - 4 overlooping species), namely 393 from Africa and Madagascar, 4,137 troglobic from Europe to Asian Russia, 84 in Middle East to Asia Central and Indian Subcontinent, 1,106 in E & SE Asia to New Guinea, 540 in Australia and Pacific Islands, and 3,303 in New World.
2 REFERENCES
[1] Gallão & Bichuette (ZooKeys, 2018), a primarily Brazilian troglobic list.
[2] White & Culver (Enciclopedia of Caves, 2019, 3th edition), general and generic diversity in caves worldwide.
[3] Freshwater Animal Diversity Assessment (Hydrobiologia, 2008), worldwide.
[4] Deharveng & Bedos (BOOK, chap. 7, 2018), worldwide.
[5] Palacios y Vargas (UMAE, 2013), for Mexico.
[6] Zampulo & Prous (Fauna Cavernicola do Brasil, 2022), for Brazil.
[7] LF Oliveira et al. (African Journal of Ecology, 2025), for Africa.
[8] J.Jugovic et al. (EJT, 2024), for Iran.
[9] Malek-Hosseini & Zamanai (Subterranean Biology, 2017), for Iran.
[10] Malek-Hosseini et al. (Subterranean Biology, 2022), for Iran.
[11]
[12] MT Guzik et al. (Invertebrate Systematics, 2010), for W Australia, unique data for this country.
[13] Díaz et al. (Biodiversidad de Cuba, cap. 15, 2015), for Cuba.
[14] Subterranean Fishes of the World (SFW) in December 15, 2024, worldwide.
[15] Furtado et al. (International Journal of Speleology, 2022) for Bolivia.
[16] Campos-Filho et al. (Nauplius, 2023) for Peru.
[17] Galán & Herrera (Boletín de la Sociedad Venezolan, Espeleologia-BSVE, 2006), for Venezuela.
[18]
[19] Angarita-Sierra (CCE/2018 | Chapter Book, 2019) + Moreno-González, J.A. et al. (American Museum Novitates, 2023) for Colombia.
[20] Chatelliers et al. (Freshwater Biology, 2009), for Clitellata worldwide.
[21] I.Turbanov et al. (PART 1, 2016 | PART 2, 2016), for former Sovietic Union.
[22] Peter Beron (Historia Naturalis Bulgarica, 2015), for SE Asia to New Guinea, including S China and S Japan.
[23] BG Nicholas (The American Midland Naturalist, 1962), for Guatemala to Panama, and Caribbean except Cuba and Jamaica.
[24] SB Peck (Canadian Journal of Zoology, 1999), for Jamaica.
[25] Culver et al. (Conservation Biology, 2021), for USA and Canada.
[26] A.Martínez et al. (BOOK, 2016), for anchialine caves in Canary Is.
[27] JL Perez-Moreno et al. (International Journal of Speleology, 2016), for anchialine caves worldwide.
[28] S.Golovatch et al. (Invertzol, 2018), for European Russia.
[29] Sket et al. (Balkan Biodiversity, 2004), for Balkans in Europe.
[30] M.Zagmajster et al. (Global Ecology and Biogeography, 2014), for European Pancrustacea.
[31] G.Pacheco et al. (Insect Conservation and Diversity, 2020), for Guatemala.
[32] Palacios-Vargas et al. (EBM, 2015), detailed checklist from Mexico.
[33] ML Niemiller et al. (Biodiversity and Conservation, 2025), detailed checklist from the USA and Canada.
[34] Hamilton Smith (Australian Journal of Entomology, 1967), for Australia.
3 WORLD CHECKLIST BY REGION
EUROPE EXCEPT RUSSIA (3,852)
Although Europe is by far the region of the world with the greatest diversity of described troglobic forms, continent-wide syntheses by group are rare, and the totals reported for many groups are often based on data from specific regions. [2] mentions 5,000 species, although the unified data compiled here does not support this number. Croatia includes 299 troglobionts and 170 stygobitic taxa, including more 100 beetles (Gottstein, S. et al., Natura Croatica, 2002).
Numbers for Europe except Russia and post-Sovietic state includes (for detailed numbers, check each group's topic):
Porifera (1, Eunapius subterraneus Sket & Velikonja, Croatia)[29],
Cnidaria (1, Velkovrhia enigmatica Matjašić & Sket, 1971, Slovenia, Croatia, and Bosnia & Hercegovina)[29],
Platyhelminthes (112, 99 Tricladida and 13 Temnocephalida)[21][29],
Nemertea (2, Prostoma puteale Beauchamp, 1932, found in France, Switzerland, and Germany, and P. hercegovinense Tarman, 1961, found in Bosnia & Herzegovina)[2],
Nematoda (7, Du Preez, Nematology, 2017, in Slovenia, Belgium, Austria, Hungary, and Romania),
Gastropoda (368, 143 stygobites from France[2], 169 stygobites from the Balkans in Falniowsk, A. et al., Hydrobiologia, 2021, and 56 troglobic[2]),
Bivalvia (1[29], Congeria kusceri Bole, 1962, Slovenia to SW Bosnia and S Dalmatia),
Annelida (103, being Marifugia cavatica Absolon & Hrabe, 1930, known from caves situated in Dinaric Karst in NW Italy, Slovenia, Croatia, and Bosnia & Hercegovina[29], two Sabelliida Nemanereis in Subterranean Biology, 2020, and 100 Clitellata[20]),
Acari (8, Belgium-4, SEE, and Italy-4, SEE | SEE),
Araneae (194, Scientific Data, 2022),
Pseudoscorpiona (132[29], Balkans),
Opiliona (21[29], Balkans),
Palpigrada (23),
Scorpiona (1, Euscorpius studentium Karaman, 2020, Montenegro, Biologia Serbica),
Symphyla (2, Slovenia),
Diplopoda (191, 29 | Iberian Peninsula | Romania),
Chilopoda (55, 50 Lithobiidae[2], Cryptopidae in Spain and Italy, and Geophilidae in France and Croatia),
Ostracoda (114[30]),
Copepoda (547[30]),
Branchiopoda (7[30], Cladocera),
Isopoda (337[30]),
Amphipoda (438[30]),
Bathynellacea (106[30]),
Thermosbaenacea (2[30]),
Mysida (3[30]),
Decapoda (16[30]),
Colembola (338, Fiera, C. et al, JZSER, 2021),
Diplura (110, Sendra et al., Zoological Journal of Linnean Society, 2021),
Coleoptera (607, 384 of them from Western Europe in Balkans[29], 25 in Spain/SEE, 54 in Ariege in France/SEE, 143 in Romania/SEE, 1 in Italy/SEE),
Dermaptera (1, Canary Islands, ZooKeys, 2019),
Zygentoma (1, W Balkans)[29],
Diptera (2, W Balkans)[29],
Hemiptera (4 from Canary Islands, Hoch et al., Subterranean Biology, 2025),
Actinopteri (2, Telestes karsticus Marčić and Mrakovčić 2011 from Croatia, and Barbatula sp. from S Germany)[14], and
Amphibia (1[29], Proteus anguinus Laurenti, 1768), totalling 3,858 troglobic in Europe.
Europe (especially Balkans) shared with Brazil and Mexico the unique troglobic Porifera worldwide (1, in Balkans), shared with Brazil the unique Bivalvia worldwide (1, in Balkans), shared with Brazil and New Zealand the unique troglobic Nemertea (2, France, Switzerland, Germany, Bosnia and Herzegovina), shared with USA the unique troglobic Amphibia (1, Italy, Slovenia, Croatia, and Bosnia-Herzegovina), shared with Colombia (possibly) the unique Nematoda worldwide (7, Belgium, Austria, Hungary, Slovenia, Romania), and has exclusivity among Temnocephalida (15), freshwater troglobic Sabellida (1, Italy, Slovenia, Croatia, Bosnia & Hercegovina) and Cnidaria worldwide (1, Slovenia, Croatia, Bosnia & Hercegovina).
Troglobic Onychophora, Symphyla, and Pauropoda are fully unknown in Europe in any form.
The Canary Islands are the richest volcanic region in the world in subterranean adapted fauna (294 troglobionts and stygobionts), followed by the Hawaiian Islands and the Undara Cave in Australia. Most of the subterranean adapted aquatic fauna from the Canary Islands is restricted to the anchialine environments in La Corona lava tube in Lanzarote, while the oligohaline stygobiont fauna, usually found in groundwater or interstitial freshwaters, is scarcer and represented by a few species of amphipods, copepods, and a single polychaete annelid recorded from Fuerteventura. In fact, Cueva de Felipe Reventón (37) and Cueva del Viento-Sobrado (36), both in Tenerife, are the second and third caves with the greatest number of troglobiont species in the world ranking, while La Corona lava tube, in Lanzarote, is the fourth richest in anchialine species (Nuñez, J., Subterranean Biology, 2020).
AFRICA AND MADAGASCAR (393)
L.F.Oliveira et al. (African Journal of Ecology, 2025)[7] lists 379 troglobic species in Africa.
PLATYHELMINTHES (4) — Tricladida (Dendrocoelum-2, Novomitchellia-1), Neorhabdocoela (1, unknown genus).
MOLLUSCA/GASTROPODA (17) — Atebbania-1, Belgrandiella-1, Bythinella-1, Giustia-7, Heideella-3, Iglica-1, Maroccopsis-1, Neohoratia-1, and Thapsia-1.
ANNELIDA (4) — Astacopsidrilus-1, Metataxis-1, Pristina-1, Trichodrilus-1.
ONYCHOPHORA (1) — Peripatopsis-1 (South Africa).
ACARI/TROMBIFIFORMES (1) — Anomalothrombium-1.
ARANEAE (16) — Acontius-1, Agraecina-1, Andoharano-4, Bathyphantes-1, Blanioonops-1, Cangoderces-1, Dysdera-2, Loxosceles-1, Meta-1, Paraleptoneta-1, Theotima-1, and Troglohyphantes-1.
SCHIZOMIDA (1) — Trithyreus-1 (Tanzania).
PSEUDOSCORPIONA (7) — Botswanoncus-1, Gymnobisium-1, Istriana-1, Neobisium-1, Roncus-2, and Selachochthonius-1.
OPILIONA (2) — Speleomontia-1 and Speleosiro-1.
AMBLYPYGI (1) — Charinus-1.
CHILOPODA (3) — Scolopendromorpha (Cryptops-1, Otostigmus-1) and Lithobiomorpha (Lithobius-1).
DIPLOPODA (4) — Dalodesmidae (Harpethrix-1), Paradoxosomatidae (Jeekelosoma-1), Chordeumatida (Origmatogona-1), and Spirostreptidea (Odontostreptus-1).
SPELAEOGRIPHACEA (1) — Spelaeogriphus-1 (South Africa).
OSTRACODA (4) — Marococandona-2, Meischcandona-1, and Namibcypris-1.
INGOLFIELLIDA (1) — Ingolfiella-1.
AMPHIPODA (81) — Afridiella-1, Austroniphargus-2, Bogidiella-4, Bollegidia-1, Crangonyx-1, Davicookia-1, Dulzura-1, Dussartiella-3, Echinogammarus-2, Eriopisella-2, Gammarus-1, Hadzia-1, Longipodacrangonyx-1, Libertinia-2, Maghrebidiella-3, Metacrangonyx-24, Nubigidiella-1, Paramellita-1, Proleleupia-1, Psammogammarus-1, Pseudoniphargus-7, Reinhardia-1, Ruffohyale-2, Salentinella-1, Sandro-2, Sternophysinx-7, Stygobarnardia-1, Trogloleleupia-5, and Tunisopisa-1.
ISOPODA (84) — Acanthastenasellus-1, Afrocerberus-1, Annina-1, Anopsilana-1, Castellanethes-6, Cirolana-1, Gabunillo-1, Haptolana-1, Johannella-1, Magniezia-6, Marocolana-1, Metastenasellus-7, Microcerberus-1, Microcharon-10, Namibianira-4, Niambia-3, Proasellus-11, Protocerberus-1, Protojanira-2, Protojaniroides-1, Saharolana-1, Skotobaena-2, Stenasellus-7, Thyphlocirolana-10, Trichoniscus-2, and Turcolana-1.
CARIDEA (13) — Caridina-7, Parisia-4, Typhlocaris-1, and Typhlopatsa-1.
THERMOSBAENACEA (5) — Monodella-1, Tethysbaena-3, and Thermosbaena-1 (Tunisia).
BATHYNELLACEA (24) — Acanthobathynella-1, Agnathobathynella-1, Cieniobathynella-1, Cteniobathynella-1, Ctenophallonella-1, Haplophallonella-1, Heterodontobathynella-1, Hexabathynella-5, Iberobathynella-1, Lamtobathynella-1, Nannobathynella-1, Nilobathynella-1, Notobathynella-1, Nunubathynella-1, Paraiberobathynella-1, Parvulobathynella-2, Racovitzaibathynella-2, and Thermobathynella-1.
BRACHYURA (2) — Skelosophusa-2 (Madagascar).
COPEPODA (67) — Copepoda/Calanoidea (Arctodiaptomus-1), Copepoda/Harpacticoida (Elaphoidella-2, Leptocaris-1, Kinnecaris-7, Monodicaris-1, Nitokra-1, Nitocrella-2, Nitocrellopsis-2, Parapseudoleptomesochra-2, Parastenocaris-20, Schizopera-2, Schizoperoides-1), and Copopoda/Cyclopoida (Allocyclops-3, Bryocyclops-4, Haplocyclops-4, Halicyclops-1, Megacyclops-1, Mesocyclops-2, Metacyclops-1, Microcyclops-2, Psammophilocyclops-2, Speocyclops-1, Thermocyclops-3, Typhlamphiascus-1).
COLLEMBOLA (4) — Acherontiella-1, Pseudosinella-1, Troglopedetes-1, and Typhlogastrura-1.
DIPLURA (3) — Anisocampa-1, Jeannelicampa-1, and Tachycampa-1.
HEMIPTERA (1) — Typhlobrixia-1.
DIPTERA (1) — Sergentomyia-1 (Tanzania).
ORTHOPTERA (1) — Spelaeiacris-1.
HYMENOPTERA (1) — Phaenopria-1 (Tanzania).
BLATTARIA (5) — Alluaudellina-1, Nocticola-3, and Typhloblattodes-1.
COLEOPTERA (22) — Antoinella-2, Apteranillus-2, Apteraphaenops-2, Anopsapterus-1, Atheta-1, Brachynillus-1, Domene-3, Duvalius-1, Ifridytes-1, Laemostenus-3, Oedichirus-1, Subilsia-1, Timosmithus-1, Trechus-1, and Typhlozyras-1.
LEPIDOPTERA (1) — Wegneria-1.
DERMAPTERA (1) — Diplatys-1.
ACTINOPTERI (1) — Clarias-1 (Namibia).
36 countries of Africa has troglobic[7], mainly in Morocco (92), Madagascar (64), Algeria (45), South Africa (34) and Namíbia (16).
Addionally, five countries in Africa has 10 troglobic Actinopteri: Cameroon (1:1/1), Somalia (2:3/3), D.R.Congo (1:1/1) and Madagascar (2:2/5)[14]; two troglobic Scorpion is cited for Madagascar — Troglotityobuthus gracilis (Fage, 1946) and Opisthacanthus pauliani Lourenço & Goodman, 2008 (Zoosystema, 2008); and [20] cites Haplotaxis gastrochaetus Yamaguchi (1953) and Villiersia guanivora Omodeo (1987), both Haplotaxidae, from Guinea — 14 added species.
Thus, Africa has 393 troglobic species. For details among Malacostraca, Copepoda and Ostracoda from Africa, see Raoul, T.K. et al (Crustaceana, 2012). Aziza cave in SE Morocco includes remarkable 22 troglobiotic and 4 stygobitic species, mainly in Coleoptera (5), Araneae (4), Entomobryomorpha (3) and Isopoda (2), by S.Moutaouakil et al. (Subterranean Biology, 2024).
FORMER USSR EXCEPT BALTIC COUNTRIES (276)
[21] lists 287 spp. from former USSR except Baltic countries (EX-USSR/SS).
RUSSIA (150, mainly in Crimea and Caucasus)
Tricladida-3, Annelida-2, Gastropoda-9, Bivalvia-1, Opiliones-5, Pseudoscorpiona-3, Acari-5, Chilopoda-1, Diplopoda-5, Ostracoda-4, Copepoda-19, Bathynellacea-6, Amphipoda-22, Isopoda-19, Decapoda-1, Collembola-21, Diplura-6, and Coleoptera-18.
ABKHAZIA (68)
Tricladida-1, Annelida-4, Gastropoda-12, Bivalvia-2, Opiliones-3, Palpigrada-1, Chilopoda-1, Diplopoda-4, Copepoda-5, Amphipoda-12, Isopoda-3, Decapoda-4, Collembola-6, and Coleoptera-10.
GEORGIA (35)
Annelida-3 (includes Erpobdella absoloni Johansson, 1913, also in W Europe), Gastropoda-4, Bivalvia-1, Opiliones-1, Pseudoscorpiona-2, Diplopoda-1, Ostracoda-1, Copepoda-7, Bathynellacea-1, Amphipoda-2, Isopoda-3, Decapoda-2, Collembola-2, Diplura-1, and Coleoptera-4.
UKRAINE (13)
Bathynellacea-3, Gastropoda-1, Amphipoda-1, Collembola-4, Coleoptera-3, and Copepoda-1.
ARMENIA (1)
Bivalvia-1.
AZERBAIJAN (4)
Amphipoda-2 and Collembola-2.
KAZAKHSTAN (2)
Amphipoda-2.
UZBEKHISTAN (5)
Copepoda-1, Annelida-1, Bathynellacea-1, Isopoda-1 and Collembola-1.
KYRGYSTAN (5)
Bathynellacea-4 and Isopoda-1.
TURKMENISTAN (4)
Isopoda-2, Gastropoda-1 and Coleoptera-1.
Belarus, Moldova and Tajikistan does no has troglobic. Five of these species are shared: two Russia/Abkhazia (Conulopolita cavatica/Gastropoda and Nemaspela abchasica/Opiliones), 1 Armenia/Georgia (Euglessa subterranea, Bivalvia), two in Russia/Abkhazia/Georgia (Bryocamptus innominatus and Echinocamptus georgevitchi, both Copepoda). After excluding 7 redundant repetitions (12 repeated entries involving these 5 species), we arrive at 280 validly cited species so far: Tricladida-4, Annelida-10, Gastropoda-26, Bivalvia-4, Opiliones-8, Palpigrada-1, Pseudoscorpiona-5, Acari-5, Chilopoda-2, Diplopoda-10, Ostracoda-5, Copepoda-29, Bathynellacea-15, Amphipoda-41, Isopoda-29, Decapoda-7, Collembola-36, Diplura-7, and Coleoptera-36.
Additionally, one more species troglobic species is listed for Kyrgystan (Chthonius kirghisicus); two more for Turkmenistan in articles: Turkmenocampa mirabilis Sendra & Stoev, 2017 (Diplura), found in Kaptarhana cave, in Lebap province (Sendra, A. et al., Subterranean Biology, 2017), and Paracobitis starostini Parin, 1983 (Actinopteri, Nemacheilidae)[14]; and at least 12 spp. (WHC Unesco) in informal lists: Gammarus troglomorphus, G. parvioculatus, and Bryocyclops jankowskajae as stygobitic Arthropoda; Kainarella minima, Pseudocaspia ljovushkini, P. starostini, P. kainarensis, and Martensamnicola cf. brevicula as stygobitic Gastropoda; and undescribed millipede Bollmania sp., a Pseudoscorpion of family Chernetidae, as well as and spiders Pholcus parthicus and Megalepthyphantes nebulosoides.
Thus, troglobic in EX-USSR/SS are 295, being Tricladida (4), Annelida (10, Erpobdella absoloni Johansson, 1913 shared with W Europa), Gastropoda (31), Bivalvia (4), Acari (5), Pseudoscorpiona (7), Araneae (2), Opiliones (8), Palpigrada (1), Chilopoda (2), Diplopoda (11), Ostracoda (5), Copepoda (30), Bathynellacea (15), Amphipoda (43), Decapoda (7), Isopoda (29), Diplura (8), Collembola (36), Coleoptera (36) and Actinopteri (1).
MIDDLE EAST EXCEPT IRAN PLUS AFGANISTAHN (13)
Middle East and Afganisthan no has unified checklist of troglobic available. At least 13 troglobic was identified for this region, in Scorpiona (1, Israel, Akrav israchanani, SEE), Diplura (1, Sendra et al., Zoological Journal of Linnean Society, 2021, in Lebanon), Annelida/Namanereis (Nereididae, in Oman-1 and Socotra-2) [2], Annelida/Clitellata (2, Oman)[20], and six Actinopteri, in Iraq (2:3/3) and Oman (1:1/3)[14].
IRAN (49)
Iran includes 51 troglobic by [8], being 46 aquatic and five terrestrial. However, we were only able to identify 48 of them in this work, in Amphipoda (24), Copepoda (12), Isopoda (3)[8], Araneae and Diplopoda one each[9], Clitellata (1, Erpobdella borisi Cichocka & Bielecki, 2015), Gastropoda (1), and (2:2/)5 Actinopteri[14]. Addtionally, one Hymenoptera was collected in latter (Yavnella laventa Griebenow, Moradmand & Isaia, 2022, Invertebrate Systematics, 2022).
INDIAN SUBCONTINENT (22)
Indian Subcontinent (Pakistan, India, Bangladesh, Nepal, Butan and Maldivas) no has unified checklist of troglobic available. 21 troglobic Actinopteri are known from this regions, in India (7:10/21)[20], and one Diplura, also in India (Sendra et al., Zoological Journal of Linnean Society, 2021).
JAPAN (90)
Japan no has unified checklist of troglobic available — nearly 200 spp. of troglobic are known from the groundwaters of Japan, most of these troglobic species, 16 of 77 genera, and what is more, 4 of 47 families are endemic to Japan. Uchidastygacaridae, Nipponacaridae, and Kantacaridae are endemic acaridan families of Japan (Matsumoto, Kôichi, International Journal of Speleology, 1976).
Two Clitellata[20], two Staphylinidae (Hlavac et al., Subterranean Biology, 2006), 80 Coleoptera in Shikoku (Ecography, 2006), and one Diplura (Sendra et al., Zoological Journal of Linnean Society, 2021) are troglobic in Japan. (1:1/)5 troglobic Actinoperi, all in Luciogobius (Gobiidae), are known from Japan[14].
KOREA (11)
11 troglobic species are recorded from the Korean Peninsula, all occurring in South Korea: Uenohadesina styx Smetana, 2000 (Staphylinidae, Hlavac et al., Subterranean Biology, 2006), seven Spelaeochthonius (Pseudotyrannochthoniidae, Plos One, 2025), and 3 Pseudocrangonyx (Pseudocrangonyctidae, PeerJ, 2021). Byung-Woo Kim et al. (Korean J. Environ. Biol., 2004) listed 257 cave-dwelling species from the Korean Peninsula — however, the study does not clearly distinguish troglobic from non-troglobic species, which makes it of limited use for assessing the region’s troglobic fauna.
CHINA (724)
We lists 742 troglobic species in China, in Annelida (2, Haplotaxida, Arhynchobdellida), Gastropoda (2, Pulmonata), Araneae (104), Pseudoscorpiona (40), Chilopoda (1), Diplopoda (114), Copepoda (23), Bathynellacea (1, Sinobathynella), Amphipoda (26), Decapoda (23), Isopoda (18), Collembola (42, mainly Coecobrya and Sinella), Diplura (1), Orthoptera (40, all Tachycines), Coleoptera (159), Hemiptera (1), Diptera (2), Actinopteri (Cypriniformes-148, Siluriformes-1) and Anura (1).
[2] cites one cave-dwelling Anura in China, however is fully rejected here as troglobic, being excludes in this count. [14] lists only 120 troglobic Actinoperi in China (data accepted here), being Amblycipitidae-1, Siluridae-1, Balitoridae-3, Cobitidae-8, Cyprinidae-38 and Nemacheilidae-69.
Additionally, five more troglobic are confirmed for China: Annelida/Hirudinea (Novataxa), and Diplura (5, more 4 spp., Sendra et al., Zoological Journal of Linnean Society, 2021).
For these numbers, China includes 716 troglobic species.
SE ASIA FROM MYANMAR TO NEW GUINEA, TIMOR AND PHILIPPINES (281)
[22] lists all the non-Actinopteri troglobionts from SE Asia to New Guinea. Their list contains inconsistencies between the manual counts and the numbers provided in the abstract, namely 15 fewer species are counted in SE Asia and 6 fewer in New Guinea. Considering only the 11 politically recognized SE Asian countries up to Papua New Guinea (Myanmar, Thailand, Laos, Vietnam, Cambodia, Malaysia, Cingapure, Indonesia, Philippines, Brunei, East Timor, and Papua New Guinea), excluding China and Japan members, we total 147 troglobic species, being
Phyllodocida (1, Namanereis beroni Hartmann-Schröder & Marinov, 1977 from PNG),
Gastropoda (1, Selmistomia beroni Bernasconi, 1995, PNG),
Araneae (13),
Pseudoscorpiones (4),
Scorpiona (2, Troglokhammouanus steineri Lourenço, 2007 in Laos and Vietbocap canhi Lourenço & Pham, 2010 in Vietnam),
Thelyphonida (1, Typopeltis magnificus Haupt, 2004, from Laos),
Opiliones (3),
Acari (4),
Diplopoda (15),
Amphipoda (3, Thailand and Sarawak, all Bogidiellidae),
Isopoda (22),
Decapoda (22),
Diplura (2),
Collembola (23),
Orthoptera (1, Eutachycines cassani (Chopard, 1954), Laos),
Blattaria (8),
Coleoptera (18),
Hymenoptera (1, Leptogenys khammouanensis Roncin & Deharveng, 2003) and
Diptera (3, both Phlebotomus from Philippines).
Additionally, are documented from this regions 37 new troglobic, namely Chaerilus cavernicola Pocock (Scorpionida, Indonesia, Invertebrate Systematics, 2008), Chaerilus chapmani Vachon & Lourenço (Scorpionida, Malaysia, Invertebrate Systematics, 2008), Orphnaecus pellitus Simon, 1892 (Theraphosidae, Philippines, Subterranean Biology, 2025), two in Diplura (Myanmar-1 and PNG-1, in Sendra et al., Zoological Journal of Linnean Society, 2021), three in Thermosbaenacea (Theosbaena, in Thailand-2 and Cambodia-1, see discussion in Thermosbaenacea topic), and 29 in Actinopteri (Myanmar-1, Thailand-4:5/9, Laos-2:3/3, Vietnan-3:6/7, Malaysia-1, Philippines-2:2/3, Indonesia-3:3/3 and Papua New Guinea-1:1/2)[14]. Two species that were supposedly identified as probable troglobic species in [2] are not recognized as such in [22]: one Temnocephalida and Leiobdella jawarerensis Richardson, 1974 (Clitellate/Haemadipsidae), both for Papua New Guinea.
Additionally, the Towakkalak and Saripa systems in Sulawesi, Indonesia (Diversity, 2021), add 32 species to this initial SE Asian dataset, including species that were mistakenly not highlighted in the text, as well as species that were not previously recognized as troglobic in the original list: Tricladida (2, Dugesia), Acari (Leeuwenhoekiidae-1), Amblypygi (Charinidae-1), Araneae (Ctenidae-1, Pholcidae-2), Opiliones (1), Palpigradi (1, Prokoenenia celebica), Pseudoscorpiones (1), Schizomida (1), Diplopoda (1, Eutrichodesmus reductus), Amphipoda (1, Bogidiellidae), Isopoda (3, Cirolanidae-1, Philosciidae-1, Armadillidae-1), Decapoda/Caridea (Atyidae-4), Decapoda/Brachyura (Gecarcinucidae-1), Collembola (Neanuridae-1, Neelidae-1, Oncopoduridae-1, Sminthuridae-1, Entomobryidae-1), Diplura: (1, Lepidocampa hypogaea), Zygentoma (Nicoletiidae-1), Blattodea (Nocticolidae-2), Coleoptera (Carabidae-1) and Hemiptera (Cixiidae-1).
In turn, Mo So Cave and the Hon Chong Karst of Vietnam (Diversity, 2023) adds 65 species to this list, being Gastropoda (Pseudoiglica-1), Acari (Leeuwenhoekiidae-1), Opilioacaridae (Siamacarus-1), Amblypygi (Weygoldtia-1), Araneae (19, Ctenidae-2, Latouchia schwendingeri Decae, 2019, Ochyroceratidae-3, Oonopidae-3, Pholcidae-4, Telemidae-5, Tetrablemmidae-1), Opiliones (3, Epedanidae-2, family undet.-1), Diplopoda (6, Haplodesmidae-1, Pyrgodesmidae-1, Trichopolydesmidae-1, Siphonophoridae-1, Cambalopsidae-2), Amphipoda (Bogidiellidae-1), Isopoda (4, family undet.-1, Sumatrillo-1, Burmoniscus-1, Stenasellus-1), Collembola (15, Hypogastruridae-2, Tullbergiidae-1, Entomobryidae-5, Isotomidae-2, Paronellidae-2, Neelidae-3), Diplura (Japygidae-1), Zygentoma (3), Blattodea (Spelaeoblatta-1), Coleoptera (3, Eustra honchongensis, Curculionidae-1, Pseudochillus honchongensis), and Heteroptera (5, Reduviidae-1, Schizopteridae-1, Cixiidae-2, Delphacidae-1, Kinnaridae-1).
After these additions and considerations, numbers by country/or region: Vietnan (89), Thailand (49), Indonesia/Sulawesi (40), New Guinea (25), Malaysian Sarawak (19), Laos (17), Philippines (16), Sumatra and adjacent Batu (6), Java and adjacent Sumba (6, including one scorpion), Indonesian Kalimatan (4), Myanmar (4), continental Malaysia (4), Cambodia (2), Malaysian Sabah (1), and Moluccas (1).
Thus, SE Asia includes 281 troglobic species.
AUSTRALIA (435)
Australia no has unified checklist of troglobic available, and the lack of precision in the numbers from Australia is absurdly complicating and confusing. Halse (On the Ecology of Australia’s Arid Zone, 2018) presents several inconsistent listings of Australia’s 'subterranean fauna', focusing on strongly positively extrapolated estimates. Some of these figures are at least comparable to the estimated or described troglobic fauna of other regions (e.g., Hemiptera, Diplopoda), whereas others appear highly unrealistic (e.g., Schizomida-59, Diplura-90, Symphyla-38, Annelida-74). These numbers are not considered here, as it is not satisfactorily clear to what extent they actually refer to truly troglobic species.
Our survey on this site, based on direct consultation of freely available online information, allowed us to compile a stable and minimally verifiable list of 435 troglobic species in Australia, within
Gastropoda (4),
Annelida/Phylodocida (1, Namanereis pilbarensis Glasby, Fiege & Van Damme, 2014, Pilbara, Zoological Journal of the Linnean Society, 2014),
Annelida/Phreodrilidae (Astacopsidrilus novus Jackson, 1931)[20],
Acari (EPA-9),
Scopiona (2, Invertebrate Systematics, 2008 | British Arachnological Society, 2001),
Palpigradi (1, Eukoenenia guzikae, Invertebrate Systematics, 2008),
Opiliones (EPA-2),
Araneae (EPA Mesa-1 | Jenolan-2 | EPA-2 | Undara-8),
Pseudoscorpiona (EPA Mesa-2 | Jenolan-1 | EPA-2 | Undara-1),
Schizomida (EPA Mesa-6 | Undara-1),
Chilopoda (4, EPA Mesa-1 | Records of the Western-2 Cryptops, 2006 | Undara-1 Scutigeridae),
Diplopoda (EPA Mesa-2 | Invertebrate Systematics-3 | Undara-4),
Symphyla (1, Hanseniella magna Scheller in Tasmania, Australian Journal of Entomology, 1996),
Ostracoda (74),
Copepoda (83),
Thermosbaenaceae (Halosbaena tulki Poore & Humphreys, 1992 is Western Australia, Wikipedia),
Anaspidacea (Jenolan-1 | EPA-1),
Bathynellaceae (1, EJT),
Amphipoda (140, Diversity, 2024),
Isopoda (EPA-2 | Undara-3),
Caridea (EPA-2, Parisia, Paratya),
Collembola (Jenolan-3 | EPA-2 | Undara-2),
Diplura (EPA Mesa-4 | Records of the Western Australian-1 | Undara-2),
Thysanura (EPA Mesa-1),
Blattodea (9, EPA Mesa-2 | EPA-5 | Undara-2),
Orthoptera (EPA-10),
Heteroptera (EPA-2),
Psocoptera (EPA-2),
Diptera (EPA-4),
Lepidoptera (EPA-1),
Coleoptera (29, EPA-27 | Journal of the Australian Entomological Society-2), and
Actinopteri (3, Synbranchidae-1, Milyeringidae-2, all from Western Australia)[14].
Halse (On the Ecology of Australia’s Arid Zone, 2018) further suggests that Western Australia may represent a potential hotspot exceeding the Balkans, currently regarded as the world’s richest hotspot of subterranean biodiversity, with estimates reaching up to 5,000 troglobionts across its vast semi-arid region. Eberhard & Howarth (Diversity, 2021) reports 19 subterranean biodiversity hotspots in the country, many of which are not included in the filtered figures presented above. If all species cited as troglobic were indeed confirmed as such, and combined with the undetermined hotspot totals—assuming each species to be restricted to a given hotspot — the resulting estimate would reach 733 spp.
In Australia, the terms 'troglofauna' and 'stygofauna' are commonly used, particularly in environmental impact assessments (EIA), to refer to all species collected from terrestrial and aquatic subterranean habitats, respectively. These terms do not distinguish ecological categories, although they are sometimes misinterpreted by non-specialists as equivalent to 'troglobiont' and 'stygobiont'. Regardless of this terminological ambiguity, precise determination of a species’ ecological status is often not critical in most Australian EIA contexts, where the primary focus is on species conservation and extinction risk. The latter is typically assessed based on the sampled and/or inferred distribution range of species in relation to the proposed mining footprint or other areas of potential impact.
Under these conditions, the characterization of Australian troglofauna becomes highly inconsistent and problematic — a truly chaos.
OCEANIA EXCEPT AUSTRALIA AND NEW GUINEA (105)
Subterranean fauna are well documented for New Zealand, where 102 described species are known from groundwater habitats (A Guide to the Groundwater Invertebrates of New Zealand, 2003): Phyllodocida (Namanereis tiriteae (Winterbourn 1969), Clitellata/Phreodrilidae (2)[20], Mollusca (10, all in Hydrobiidae), Acari/Hydracarina (71), Bathynellaceae (5), Anaspidaecea (2, details in Stygocarella pleotelson Schminke, 1980 and Stygocaris townsendi Morimoto, 1977, Beaufortia, 1980), Amphipoda (4), Isopoda (4), and Coleoptera (3).
Additionally, at least one species of troglobic Nemertea is known in New Zealand, Exosphaeroides quirosi Jaume et Queinnec, 2007 (Isopoda) for Vanuatu[22], and one Actinopteri from Guam (Eleotris undescribed species)[14].
Stewart, Robertson & Kasper (NZ Speleological Bulletin, 2023) discuss cave fauna in New Zealand in a highly allegorical manner, which makes the data difficult to use and, therefore, not considered here.
USA/CANADA (1,476)
[33] lists all troglobic from the USA and Canada, which includes Alaska and Hawaii, with (124:288/)1,461 spp., representing six phyla, namely Platyhelminthes (6:7/34), Syndermata (1:1/1), Annelida (4:8/22), Gastropoda (6:17/39), Nematoda (2:2/2), Acari (13:21/32), Araneae (8:23/121), Opiliones (7:12/46), Pseudoscorpiones (8:27/153), Schizomida (2:2/2), Scorpiones (1:1/1), Copepoda (5:7/18), Ostracoda (2:14/22), Bathynellacea (2:7/23), Thermosbaenacea (1:1/1), Amphipoda (11:24/192), Isopoda (7:21/118), Decapoda (4:7/44), Chilopoda (2:4/4), Diplopoda (13:27/140), Collembola (7:16/101), Diplura (2:6/10), Coleoptera (9:31/284), Dermaptera (1:1/1), Diptera (3:3/3), Hemiptera (3:4/10), Lepidoptera (1:1/1), Orthoptera (1:2/6), Zygentoma (1:2/8), Actinopterygii (3:7/10) and Amphibia (1:2/12).
Additionally, Salishbaena kootenai Wagner & Reid, 2021 (Thermosbaenacea, Crustaceana, 2021), two Diplura (EJT, 2026) and 12 more Actinoperi[14] are known as troglobic from region. Thus, USA (including Hawaii and Alaska) and Canada includes 1,476 troglobic, as Platyhelminthes (6:7/34), Syndermata (1:1/1), Annelida (4:8/22), Gastropoda (6:17/39), Nematoda (2:2/2), Acari (13:21/32), Araneae (8:23/121), Opiliones (7:12/46), Pseudoscorpiones (8:27/153), Schizomida (2:2/2), Scorpiones (1:1/1), Copepoda (5:7/18), Ostracoda (2:14/22), Bathynellacea (2:7/23), Thermosbaenacea (2:2/2), Amphipoda (11:24/192), Isopoda (7:21/118), Decapoda (4:7/44), Chilopoda (2:4/4), Diplopoda (13:27/140), Collembola (7:16/101), Diplura (2:6/12), Coleoptera (9:31/284), Dermaptera (1:1/1), Diptera (3:3/3), Hemiptera (3:4/10), Lepidoptera (1:1/1), Orthoptera (1:2/6), Zygentoma (1:2/8), Actinopterygii (3:7/22) and Amphibia (1:2/12).
However, it is important to note that the list for this region includes anchialine forms from Hawaii and parasitic forms in Nematoda, Platyhelminthes, and Acanthocephala — groups that are controversial and/or outside the scope of this text, but which are retained here for the sake of numerical consistency.
Some genera represent remarkable subterranean radiations in the USA, such as Pseudophthalmus (147, Coleoptera), Stygobromus (139, Amphipoda), Pseudotremia (62, Diplopoda), Caecidotea (56, Isopoda), Cicurina (52, Araneae), Tyrannochthonius (36, Pseudoscorpiona), Pygmarrhopalites (31, Collembola), and Batrisodes (30, Coleoptera). Together, these 8 genera account for 553 troglobic species in the USA, representing nearly 2/5 of the national diversity.
Only 11 spp. occur as troglobic in Canada, being one Rhagidiidae, three Araneae (Linyphiidae), one Collembola/Entomobryidae, one Collembola/Arrhopalitidae, four Amphipoda/Stygobromus, and one Isopoda/Asellidae. In Hawaii, there are 60: two Rhagidiidae, five Araneae (two Linyphiidae, two Lycosidae, one Ochyroceratidae), five Pseudoscorpiones/Chthoniidae, five Collembola/Entomobryidae, one Collembola/Neanuridae, four Coleoptera/Carabidae, two Diptera, and all troglobic Hemiptera (10), Dermaptera (1), Lepidoptera (1), and Orthoptera (6) from the USA, ten Amphipoda, four Isopoda/Philosciidae, two Atyidae, and two Procarididae. Only two troglobic occur in Alaska: one Amphipoda and one Bathynellacea.
Excluding Canada, USA (with Hawaii and Alaska) includes Platyhelminthes (6:7/34), Syndermata (1:1/1), Annelida (4:8/22), Gastropoda (6:17/39), Nematoda (2:2/2), Acari (13:21/31), Araneae (8:23/118), Opiliones (7:12/46), Pseudoscorpiones (8:27/153), Schizomida (2:2/2), Scorpiones (1:1/1), Copepoda (5:7/18), Ostracoda (2:14/22), Bathynellacea (2:7/23), Thermosbaenacea (2:2/2), Amphipoda (11:24/188), Isopoda (7:21/117), Decapoda (4:7/44), Chilopoda (2:4/4), Diplopoda (13:27/140), Collembola (7:16/99), Diplura (2:6/12), Coleoptera (9:31/284), Dermaptera (1:1/1), Diptera (3:3/3), Hemiptera (3:4/10), Lepidoptera (1:1/1), Orthoptera (1:2/6), Zygentoma (1:2/8), Actinopterygii (3:7/22) and Amphibia (1:2/12).
BRAZIL/USA COMPARATION
Neither Brazil nor the USA have troglobic in Cnidaria. Troglobic lineages in Brazil unknown as troglobic in USA are Porifera (2 ✕ 0), Nemertea (3 ✕ 0), Onychophora (1 ✕ 0), Bivalvia (1 ✕ 0), Pauropoda (1 ✕ 0), Symphyla (14 ✕ 0), Blattodea (8 ✕ 0) and Hymenoptera (3 ✕ 0).
Troglobic lineages in USA unknown as troglobic in Brazil include eight, 5 parasitic worms, namely Platyhelminthes Alloeocoela (1/1), Cestoda (1/1) and Trematoda (1/1), Syndermata/Acanthocephala (1/1), Nematoda (2/2), Thermosbaenacea (1/1), Lepidoptera (1/1), and Amphibia (1:2/12).
At commom groups, Brazil and USA are fully tied in Dermaptera,Orthoptera, and Sarcoptiformes.
At commom groups, USA surpasses the Brazil (US excedents in parentesis) in
Coleoptera (3F-16G-192S), Amphipoda (6F-19G-158S), Pseudoscorpiones (2F-20G-90S), Diplopoda (5G-42S), Decapoda (3F-6G-37S), Araneae (30S), Bathynellacea (1F-6G-22S), Trombidiformes (4F-12G-21S), Ostracoda (3F-13G-21S), Copepoda (3F-5G-14S), Platyhelminthes/Tricladida (13S), Annelida (2F-6F-13S), Gastropoda (10G-12S), Opiliones (4G-12S), Diplura (3G-3S), Mesostigmata (1F-1G-2S), Diptera (2F-2G-2S), Schizomida (1F-1G-1S) and Collembola (4G).
At commom groups, Brazil surpasses ot tied the USA in at all levels in Chilopoda, Scorpiona, Palpigrada, Zygentoma, Hemiptera, Isopoda and Actinopteri.
MEXICO (523)
[5] lists 524 troglobic species in Mexico:
Platyhelminthes (5, Dugesia-3, Opisthobursa-2),
Nematoda (1, Axonchium sbordonii Zullini, 1974),
Annelida/Nereididae (1, Nemanereis cavernicola Solis-Weiss & Espinasa, 1991), Haplotaxida/Acanthodrilidae (Eodrilus-2), Branchiobdellida-Branchiobdellidae (Cambarincola-2),
Gastropoda (8, Amnicolidae-2, Hydrobiidae-3, Lithoglyphidae-3),
Arachnidae/Scorpiones (16, Diplocentridae-4, Euscorpiidae-2, Typhlochactidae-7, Vaejovidae-3),
Arachnidae/Pseudoscorpiona (40, Bochicidae-18, Cheliferidae-1, Chthoniidae-12, Ideoroncidae-7, Syarinidae-2),
Arachnidae/ Schizomida (23, Hubbardidae-14, Protoschizomidae-9),
Arachnidae/Amblypygi (Paraphrynus-6),
Arachnidae/Araneae (58, Agelenidae-2, Cicurinidae-4, Dipluridae-3, Leptonetidae-6, Linyphiidae-1, Nesticidae-4, Ochyroceratidae-2, Oonopidae-1, Pholcidae-25, Tetrablemmidae-1, Theraphosidae-9),
Arachnidae/Opiliones (16, Neogoveidae-1, Phalangodidae-3, Stygnopsidae-11, Nemastomatidae-1),
Arachnidae/Ricinulei (Pseudocellus-11),
Arachnidae/Acari (Prostigmata-2),
Chilopoda (Lithobiidae-2, Scolopocryptopidae-3),
Diplopoda (73, Glomeridesmida-1, Glomerida-6, Spirobolida-1, Spirostreptida-7, Chordeumatida-13, Polydesmida-45),
Copepoda (12, Harpacticoida-1, Calanoida-3, Cyclopoida-8),
Ostracoda (4, Halocyprida-2, Podocopida-2),
Malacostraca/Themosbaenaceae (Tulumella-1),
Malacostraca/Lepidomysidae (Spelaeomysis-3),
Malacostraca/Mysidae (Antromysis-2),
Malacostraca/Stygiomysidae (Stygiomysis-2),
Malacostraca/Amphipoda (22, Bogidiellidae-10, Hadziidae-12),
Malacostraca/Isopoda (52, Anthuridea-2, Flabellifera-23, Microcerberidea-1, Asellota-13, Oniscoidea-13),
Malacostraca/Decapoda (Caridea-21, Procarididea-1, Astacidea-5, Brachyura-5),
Remipedia (Xibalbanus-2),
Collembola (Poduromorpha-11, Entomobryomorpha-27, Symphypleona-4),
Diplura (Campodeidae-4),
Insecta/Zygentoma (Nicoletiidae-13),
Insecta/Orthoptera (Phalangopsidae-6),
Insecta/Hemiptera (Cixiidae-2, Kinnariidae-1),
Insecta/Coleoptera (44, Dytiscidae-1, Carabidae-30, Histeridae-4, Leiodidae-8, Ptinidae-1),
Echinodermata/Asteroidea (1, Copidaster cavernicola Solís-Marín & LaguardaFigueras, 2010), and
Actinopteri (Characidae-1, Ictaluridae-2, Heptapteridae-4, Bythitiidae-1, Synbranchidae-1).
As anchialine species in Mexico in Yucatan peninsula restricted to marine layer of these caves, 16 were excluded from this list: two Remipedia (Xibalbanus, now 3), two Copepoda (Balinella-1, Exumella-1), two Ostracoda (both Halocryptida), two Amphipoda (Bahadzia-2), one Isopoda (Metacirolana mayana (Bowman, 1987)), Procaris mexicana Sternberg & Schotte, 2004, 5 Caridea (Typhlatya dzilamensis Álvarez et al., 2005, Yagerocaris cozumel Kensley, 1988, Triacanthoneus akumalensis Álvarez et al., 2012, Callismata nohochi EscobarBriones et al., 1997, Agostocaris bozanici Kensley, 1988), and the echinoderm Copidaster cavernicola Solís-Marín & LaguardaFigueras, 2010. Additonally, nine new troglobic are adde in checklist of Mexico: one Porifera (1, Legendre et al., Subterranean Biology, 2023, undescribed), Tonalteca spinooculorum (Opiliones, ), five Rhamdia (Heptapteridae, SFW), Caecieleotris morrisi Walsh and Chakrabarty 2016 (Eleotrididae, SFW), and Lucifuga undescribed species 6 (Bythitiidae, SFW). Hlavac et al. (Subterranean Biology, 2006) cite Stenophloeus reddelli Herman, 1969 as troglobic in Mexico; however, it is rejected here[5]. With these changes, the updated list accepted here is as follows, comprising 518 spp.
Porifera (1),
Platyhelminthes (5, Dugesia-3, Opisthobursa-2),
Nematoda (1, Axonchium sbordonii Zullini, 1974),
Annelida/Nereididae (1, Nemanereis cavernicola Solis-Weiss & Espinasa, 1991), Haplotaxida/Acanthodrilidae (Eodrilus-2), Branchiobdellida-Branchiobdellidae (Cambarincola-2),
Gastropoda (8, Amnicolidae-2, Hydrobiidae-3, Lithoglyphidae-3),
Arachnidae/Scorpiones (16, Diplocentridae-4, Euscorpiidae-2, Typhlochactidae-7, Vaejovidae-3),
Arachnidae/Pseudoscorpiona (40, Bochicidae-18, Cheliferidae-1, Chthoniidae-12, Ideoroncidae-7, Syarinidae-2),
Arachnidae/ Schizomida (23, Hubbardidae-14, Protoschizomidae-9),
Arachnidae/Amblypygi (Paraphrynus-6),
Arachnidae/Araneae (58, Agelenidae-2, Cicurinidae-4, Dipluridae-3, Leptonetidae-6, Linyphiidae-1, Nesticidae-4, Ochyroceratidae-2, Oonopidae-1, Pholcidae-25, Tetrablemmidae-1, Theraphosidae-9),
Arachnidae/Opiliones (16, Neogoveidae-1, Phalangodidae-3, Stygnopsidae-12, Nemastomatidae-1),
Arachnidae/Ricinulei (Pseudocellus-11),
Arachnidae/Acari (Prostigmata-2),
Chilopoda (Lithobiidae-2, Scolopocryptopidae-3),
Diplopoda (73, Glomeridesmida-1, Glomerida-6, Spirobolida-1, Spirostreptida-7, Chordeumatida-13, Polydesmida-45),
Copepoda (10, Harpacticoida-1, Calanoida-1, Cyclopoida-8),
Ostracoda (Podocopida-2),
Malacostraca/Themosbaenaceae (Tulumella-1),
Malacostraca/Lepidomysidae (Spelaeomysis-3),
Malacostraca/Mysidae (Antromysis-2),
Malacostraca/Stygiomysidae (Stygiomysis-2),
Malacostraca/Amphipoda (20, Bogidiellidae-10, Hadziidae-11),
Malacostraca/Isopoda (52, Anthuridea-2, Flabellifera-22, Microcerberidea-1, Asellota-13, Oniscoidea-13),
Malacostraca/Decapoda (Caridea-16, Astacidea-5, Brachyura-5),
Remipedia (Xibalbanus-2),
Collembola (Poduromorpha-11, Entomobryomorpha-27, Symphypleona-4),
Diplura (Campodeidae-4),
Insecta/Zygentoma (Nicoletiidae-13),
Insecta/Orthoptera (Phalangopsidae-6),
Insecta/Hemiptera (Cixiidae-2, Kinnariidae-1),
Insecta/Coleoptera (44, Dytiscidae-1, Carabidae-30, Histeridae-4, Leiodidae-8, Ptinidae-1),
Echinodermata/Asteroidea (1, Copidaster cavernicola Solís-Marín & LaguardaFigueras, 2010), and
Actinopteri (11, Characidae-1, Ictaluridae-2, Heptapteridae-9, Bythitiidae-2, Eleotrididae-1, Synbranchidae-1).
Mexico no has described troglobic in Nemertea, Onychophora, Mollusca/Bivalvia, Bathynellacea, Spelaeogriphacea, Blattodea, Dermaptera, Diptera, Hymenoptera, Pauropoda, Symphyla, Palpigradi or Amphibia. When considering species not yet described in Brazil, the country surpasses Mexico in all groups except Nematoda (unknown in Brazil), Schizomida, Ricinulei (unknown in Brazil), Scorpionida, Mysids overall (unknown in Brazil), Ostracoda, Copepoda, Decapoda, and Zygentoma.
AMERICA CENTRAL FROM GUATEMALA TO PANAMA (21)
The only consistent reference on the troglobic fauna of southern Mexico, America Central, and the Caribbean is [23], who lists 170 spp. for the vast region. Currently, Mexico, Cuba, and Jamaica have more recent references and are treated independently. Therefore, we use these references only for the region from Guatemala to Panama and the West Indies, excluding Cuba and Jamaica.
Based on [23], only eight troglobic occur from Guatemala to Panama, six valid nowdays (exclusion of Pselliodes harveyi Chamberlin, 1942 from Panama and Pselliophora cavincola Chamberlin, 1918 from Trinidad, both under Sphendononema guildingii Newport, 1845, Chilobase), namely:
Trombicula cavernarum Ewing, 1933 and
T. trifurca Ewing, 1933 (Trombidiformes, Panama),
Otostigmus cooperi Chamberlin, 1942 (Chilopoda, Panama),
Calymmodesmus inquinatus Causey, 1960 (Diplopoda, Guatemala),
Lepidocyrtus usitatus Folsom, 1927 (Collembola, Panama), and
Ptomaphagus giaquintoi Jeannel, 1936 (Coleoptera, Alta Verapaz, Guatemala).
Additonally, 15 troglobic are added in region, including three crabs in the genus Typhlopseudothelphusa (Pseudothelphusidae, two in Guatemala and one in Belize, Hobbs, Texas Memorial Museum Speleological Monographs, 1986), two Diplura in Guatemala (Sendra et al., EJT, 2026), Rhamdia typhla Greenfield, Greenfield, and Woods, 1982 (Actinopteri/Heptapteridae, Belize)[14], and nine from Gruta de Lanquín in Guatemala[31] (Isopoda-2, Platyhelminthes-2, Araneae-1, Pseudoscorpiones-1, Opiliones-1, Polydesmida-1, and Zygentoma-1).
For these numbers, America Central has 21 documented troglobic. S.Deleva et al. (Subterranean Biology, 2023) discusses the cavernicolous fauna of Costa Rica and confirms that there are no troglobic species in the country, only strong candidates are Pseudothelphusa puntarenas Hobbs 1991 (Decapoda), various specimens of Trogolaphysa (Collembola) and some Acari (Rhagidiidae) from Barra Honda.
CUBA (130)
[13] cites 112 troglobic in Cuba, all Arthropoda: Acari (12), Scorpiones (1), Pseudoscorpiones (3), Amblypygi (1), Schizomida (5), Ricinulei (4), Opiliones (4), Araneae (7), Chilopoda (2), Ostracoda (5), Copepoda (18), Mysidacea (5), Isopoda (7), Decapoda (13), Amphipoda (3), Remipedia (1), Zygentoma (3), Collembola (7), Orthoptera (4), Blattoidea (1) and Coleoptera (6) — here Remipedia (1) is excluded due to anchialine status.
Additionally 20 species are reported as troglobic in Cuba:
Eukoenenia orghidani (Palpigradi, The Journal of Arachnology, 2010),
Alayotityus delacruzi Armas, 1973, Tityopsis sheylae Teruel & Rodríguez-Cabrera, 2020 (Scorpionida/Buthidae, Euscorpius, 2021),
Tethysbaena vinabayesi Wagner, 1994 (Thermosbaenacea, see discussion in Thermosbaenacea topic),
Clitellio cavernicolus Botea, 1983 (Tubificidae)[20], four species of Pristina (Tubificidae)[20],
two Cryptops (Chilopoda, ZooKeys), and
nine Lucifuga (Bythitidae, Ophidiiformes, Actinopteri)[14].
Thus, Cuba has 131 troglobic species in our survey.
JAMAICA (42)
Troglobic in Jamaica contains 43 spp. troglobic[24], being 27 of troglobic in Onychophora (Speleoperipatus spelaeus Peck, 1975), Araneae (9), Opiliones (2), Pseudoscorpionida (4), Schizomida (1), Isopoda (1), Collembola (2), Blattodea (1), Hemiptera (2) and Coleoptera (4), and 16 stygobitic, in Tricladida (1), Ostracoda (3), Mysida (2), Thermosbaenacea (1), Decapoda (2), Amphipoda (4) and Isopoda (3).
Thermosbaenacea record is excluded here due to inconclusive status relative to the literature — thus, we accepted 42 troglobic in Jamaica.
CARIBBEAN EXCEPT CUBA AND JAMAICA PLUS BERMUDA (27)
The only consistent reference on the troglobic fauna of southern Mexico, America Central, and the Caribbean is [23], who lists 170 spp. for the vast region. Currently, Mexico, Cuba, and Jamaica have more recent references and are treated independently. Therefore, we use these references only for the region from Guatemala to Panama and the West Indies, excluding Cuba and Jamaica.
[23] lists nine troglobic for this area, namely:
Maymena bruneti Gertsch, 1960 (Araneae, Trinidad),
Bogidiella bredini Shoemaker, 1959 (Amphipoda, Barbuda),
Metaniphargus beattyi Shoemaker, 1942 (Amphipoda, Virgin Islands),
M. nicholsoni Shoemaker, 1959 (Amphipoda, Barbuda),
Typhlatya monae Chace, 1954 (Decapoda Caridea, Puerto Rico),
Stygiomysis holthuisi Gordon, 1960 (Mysida, Saint Marteen),
Calycuoniscus spinosus Collinge, 1917 (Isopoda, Trinidad),
Porcellionides minutissimus Van Name, 1936 (Isopoda, Bahamas), and
Philoscia bermudensis Dahl, 1892 (Isopoda, Walsingham Cave, Bermuda).
Additionally, 13 more troglobic are confirmed for this area:
Pseudochthonius arubensis Wagenaar-Hummelinck (Aruba, Zoologia, 2023),
Heteronebo clareae Armas, 2001 in Bahamas (Scorpiona, Euscorpius, 2021),
Cazierius ciguayo Teruel, Jiménez & Santos, 2021 and C. cayacoa Teruel, Jiménez & Santos, 2021 in Dominican Republic (Scorpiona, Euscorpius, 2021),
Newportia stoevi Schileyko, 2013 (Chilopoda, Puerto Rico, ZooKeys, 2013),
six Namanereis (Annelida, Nereididae, one shared with Mexico),
Phallodriloides macmasterae Erséus, 1986 (Tubificae) in Bermuda[20], Dero haitiensis Dumnicka, 1986 and Spirospermoides stocki Dumnicka, 1983 (both Tubificidae) in Haiti and adjacent islands[20],
Tethysbaena colubrae (Thermosbaenacea, Puerto Rico),
Rhamdia urichi Mees, 1974 (Heptapteridae, from Trinidad and Tobago)[14], amd two Lucifuga (Bythitidae, Bahamas)[14].
Thus, in West Indies except Jamaica and Cuba are known 22 troglobics.
GUIANAS (0)
[11] cites possibly Lepidocampa (Diplura), Eukoenenia (Palpigrada) and Modisimus (Araneae) as troglobic in French Guiana, however their troglobic status in unknown, and all are rejected here.
VENEZUELA (47)
[17] lists 46 troglobic in Venezuela, in Acari (4:9), Opiliones (4, all in Trinella, Agoristenidae), Amblypygi (2, both Charinus), Scorpionida (1, Taurepania trezzii, Chactidae), Diplopoda (1, Trichopolydesmidae), Collembola (1, Onychiuridae), Copepoda/Cyclopidae (1), Ostracoda (2, in Cyprididae), Bathynellaceae (1, Parabathynellidae), six Isopoda (6:6), Amphipoda (3:3), Decapoda (2, Pseudothelphusidae), Orthoptera (2, both Rhaphidiophoridae), Blattodea (1, Paranocticolla venezuelana), Coleoptera (3:3), and seven fishes (in 4 families).
From this list, we excluded the two anchialine species — Stygocyathura univam (Botosaneanu, 1983)(Isopoda) and Metaniphargus venezuelanus (Stock & Botosaneanu, 1983)(Amphipoda), as they fall outside the scope of this article.
Additionaly, troglobic in Venezuela includes Speleopsocus chimanta Lienhard, 2010 (Psocoptera, Revue suisse de Zoologie, 2015), Krenedrilus papillatus Dumnicka, 1983 (Annelida/Tubificidae)[20], and Cryptocellus armasi Botero-Trujillo and Prendini, 2025 (Ricinulei, Zootaxa, 2025).
Thus, 47 troglobic are recorded in Venezuela.
COLOMBIA (42)
65 troglobic are listed in Colombia[19], in Nematoda (1, Rhabditida), Annelida (2, Haplotaxide, Enchytraeidae one each), Tityus grottoedensis (Scorpionida), Opiliones (7:7), Amblypygi (2, Phrynidae and Paracharontidae), Araneae (16, in 11 families, Barychelidae, Araneidae, Anyphaenidae, Lycosidae, Tetragnathidae, Pisauridae without troglobic in Brazil), Chilopoda (3, undescribed Scolocryptops and 2 Psilliodes), Diplopoda (7), Isopoda (1, Armadillidium), Coleoptera (5, two in Staphylinidae; Carabidae, Nitidulidae, Passalidae one each), Ephemeroptera (2, if confirmed, will be the first troglobis in this order worldwide), Blattoidea (1, Blaberidae), Dermaptera (1), Diptera (2, Drosophilidae and Mycetoohilidae one each), Hemiptera (3, Cydnidae, Cavernicola pilosa in Reduviidae, Strudivelia cinctipes in Veliidae), Lepidoptera (1), Trichoptera (2, in Hydrobiosidae and Hydropsychidae), Actinopteri (7, all Trichomycterus) and Avialia (Steatornis caripenis).
Steatornis caripenis assignated in work as troglobic is a obviously a huge error. Troglobic status rejecteds for both Annelida (Angarita-Sierra et al, Chapter Book, 2019), all Psilliodes (fully rejected by many sources), 13 Araneae (Angarita-Sierra et al, Chapter Book, 2019), Tityus grottoedensis (by original description, Zootaxa, 2014), 4 Opiliones (Angarita-Sierra et al, Chapter Book, 2019), one Ephemeroptera (Angarita-Sierra et al, Chapter Book, 2019), Cavernicola pilosa (Dissertation/FOC) and Strudvelia cinctipes (Biota Colombiana).
Additionally, 10 troglobic Actinopteri are recorded in Colombia (all in Trichomycterus)[14], and Diploexochus cacique (Isopoda) was described in 2025 (ZooKeys).
With these exceptions, Colombia has, with some consistency, 42 spp. of troglobic: Nematoda (1, Rhabditida), Opiliones (3), Amblypygi (2, Phrynidae and Paracharontidae), Araneae (3), Chilopoda (1, undescribed Scolocryptops), Diplopoda (6, in Chelodesmidae, Pyrodesmidae, Spirostreptidae, Stemmiulidae and Rhinocricidae), Isopoda (2, both in Armadillidae), Coleoptera (5, two in Staphylinidae; Carabidae, Nitidulidae, Passalidae one each), Ephemeroptera (1), Blattoidea (1, Blaberidae), Dermaptera (1), Diptera (2, Drosophilidae and Mycetoohilidae one each), Hemiptera (1), Lepidoptera (1), Trichoptera (2, in Hydrobiosidae and Hydropsychidae) and Actinopteri (10, all Trichomycterus) — however, troglobic status of Nematoda, Ephemeroptera, Dermaptera, both Diptera, Lepidoptera and both Trichoptera need strong validation to be fully credited.
ECUADOR (8)
Ecuador no has unified checklist of troglobic available. Literaure lists 8 troglobic species in country: Marionina ecuadoriensis Righi and Hemienchytraeus mauriliae Righi (Clitellata/Enchytraeidae)[20], also Righi (Papéis A vulsos de Zoologia, 1981), Tayos ashmolei Reddell & Cokendolpher, 1984 (Schizomida, British Arachnological Society, 1984) and Troglotayosicus vachoni Lourenço, 1981 (Scorpiona/Troglotayosicidae) from Cueva de los Tayos, Santiago-Morona Province; Ischnoptera peckorum Roth, 1988 (Ectobiidae, Blattaria, Galapagos Species Database, Bellavista Cave) and Pinostygus galapagoensis Cambell & Peck, 1989 (Staphylinidae, Hlavac et al., Subterranean Biology, 2006) from Galapagos islands; and two Actinopteri, Astroblepus pholeter Collette, 1962, and one Chaetostoma undescribed species[14].
PERU (3)
Only three spp. in Peru the following can be considered troglobic[16]: Acheroxenylla lipsae Palacios-Vargas, 2020 (Collembola, Poduromorpha, Hypogastruridae), Astroblepus riberae Cardona and Guerao, 1994 (Actinopteri, Siluriformes, Astroblepidae) and Caecopactes chullachaqui Campos-Filho, Sfenthourakis and Bichuette, 2024 (Isopoda, Scleropactidae).
BOLIVIA (8)
Bolivia no has unified checklist of troglobic available. Six troglobic are cited for Bolivia sampled in Umajalanta cave[15]: Coleoptera (3, one described, Escolatrechus bolivianus Mateu, 2002), Acari (1, Rhagidiidae), Collembola (1, Entomobryomorpha), and Isopoda (1, Dubioniscidae). Additionally, Trichomycterus chaberti Durand 1968 and Phreatobius sanguijuela Fernandez, Saucedo, Carvajal and Schaefer 2007 are troglobic in Bolivia[14], totalling 8 troglobic species in country.
ARGENTINA (31)
Without a unified checklist of troglobic available, our survey counted 31 troglobic in Argentina:
■ 24 known in Cuchillo Curá cave system, largest known cave system in country (ICS/vol VII, 2025), being 21 terrestrial and three aquatic, all Arthropoda, in Acari (1), Pseudoscorpiona/Chthoniidae (1), Opiliones/Laniatores (1), Palpigradi (2), Araneae (3), Symphyla (1), Isopoda (4), Amphipoda (2), Collembola (3), Diplura (3, Campodeidae, Japygidae), Coleoptera/Leiodidae (1), Auchenorrhyncha/Cixiidae (1), and Zygentoma/Nicoletiidae (1) — only six of these species are described: Picunchenops spelaeus Maury, 1988 (Opiliones/Laniatores), Pseudothiphistes pallidus (Millidge, 1991) - a Araneae/Linyphiidae, Notolathrus sensitivus Remes Lenicov, 1992 (Cixiidae), Adelopsis picunche Gnaspini, 1991 (Coleoptera/Leiodidae), Grossogidiella mauryi (Grosso & Fernández, 1993) and Patagongidiella danieli Grosso & Fernández, 1993), both Amphipoda/Bogidiellidae.
■7 in other caves: Troglospinotheca (1, Collembola, Neuquen region, Mémoires de Biospéologie, 1999), four Anaspidacea (Plazi | Plazi), Pleonaraius spelaeus (Diplopoda/Polydesmida, Neuquen, Romero-Rincon & Ferreira, Arthropoda, 2025), and one Actinopteri, Silvinichthys bortayro Fernandez and De Pinna 2005, in Trichomycteridae[14].
PARAGUAY, URUGUAY AND CHILE (2)
For Paraguay, Uruguay and Chile, only two troglobic are documented: two Anaspidacea (Plazi | Plazi), both from Chile.