1 ABSTRACT
In this post, we discuss the diversity of obligate troglobitic and stygobitic animals in Brazil, collectively referred to here by the term troglobic (see discussion below). A comprehensive literature review was conducted, with a strong focus on the global context, and as of January 5, 2025, 678 troglobic species have been recorded in Brazil and approx. 10,174 in World. Brazilian diversity span over 8 of the 10 Metazoa elegible phyla for troglobics: Porifera (2), Platyhelminthes (15), Nemertea (2), Annelida (6), Mollusca (22), Onychophora (1), Arthropoda (577) and Craniata/Actinopteri (27). Cnidaria and Nematoda does not has troglobics in Brazil and has described troglobics only in Europe. Each species is presented in own card, featuring distribution map and images, when available. We also conducted cross-sectional analyses, examining the literature on the distribution of these species in various regions of the world, as well as their occurrence by state, municipality, and taxonomic group in Brazil. For nomenclature, we use the notation ( z : y / ) x, which refers to x species in y genera of z families.
ALL TEN PHYLLA IN METAZOA WITH TROGLOBITICS IN METAZOA PHYLOGENETIC TREE, AFTER LAUMER C.E. ET AL. (SEE, 2023)
2 INTRODUCTION
For the purposes of this text, based on Trajano & Bichuette (BOOK, 2006), we will use the following terminology for Metazoarian in caves: (1) troglobionts - obligatory subterranean, exclusively subterranean source populations; sink populations may be found in surface habitats; if obligate subterranean aquatic, we will call stygobionts; (2) troglophiles are subterranean source populations that maintain genetic connection with the epigean ones through individuals that disperse between these habitats; (3) trogloxenes are individuals regularly found in subterranean habitats but which must return periodically to the surface to complete their life cycle; they would be instances of source populations in epigean habitats, but using subterranean resources (the obligatory trogloxenes, all individuals are dependent on both subterranean and surface resources).
These terms sometimes overlap and blend, and in many contexts, they end up being inefficient in communication between different sources. Aiming to standardize the language used in this post, we have adopted the following terms to represent the different groups of obligatorily subterranean animals:
troglobic = obligatory subterranean, with exclusively subterranean source populations; sink populations may be found in surface habitats.
troglobiont = terrestrial troglobics, used in the text if necessary.
stygobitic = freshwater or amphibious troglobics, used in the text if necessary.
halotroglobic = marine troglobics, used in the text if necessary.
Troglobics are usually recognized by the presence of troglomorphisms - autapomorphies related to the subterranan selective regime. The commonest and most conspicuous troglomorphisms are reduction up to complete loss of visual structures and dark tegumentar pigmentation; other frequent troglomorphisms include elongation of barbels in fish and of appendices in arthropods, and reduction of wings in insects.
3 SOME REFLECTIONS
When discussing the concept of troglobics, four key questions arise for analysis. The first concerns whether this concept also applies to fungi and other organisms, such as protists, which are traditionally classified separately. The second question is defining which habitats qualify a potential troglobic as a true troglobic: anchialine caves, littoral interstitial habitats, deep phreatic and hyporheic of rivers, MSS (milieu souterrain superficiel, underground network of empty air-filled voids and cracks developing within multiple layers of rock fragments, generally covered by topsoil, SEE), epikarts and hypotelminorheic habitats can be? The third issue involves defining troglobic within well-established biological groups, exploring apomorphies (derived characteristics) and the phenomenon of troglomorphism, which refers to adaptations specific to subterranean environments. Finally, the fourth point questions to what extent the concept of troglobic can be applied to marine organisms, considering the unique features of aquatic environments and their caves. Culver, Deharveng, Bedos & Pipan (Diversity, 2021) makes interesting observations about some of these points.
4 REFERENCES
[1] Gallão & Bichuette (ZooKeys, 2018) - Brazilian primarily 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).
[4] Deharveng & Bedos (BOOK, chap. 7, 2018).
[5] Palacios y Vargas (UMAE, 2013), for Mexican numbers.
[6] Zampulo & Prous (Fauna Cavernicola do Brasil, 2022).
[7] Oliveira, L.F. (THESIS, 2024), for Africa.
[8] Jugovic, J. 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] Almanaque Z/South America Cave Smaller Divesities, for Guyana, Suriname, French Guiana, Argentina, Chile, Paraguay and Uruguay.
[12] M. T. 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.
[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] Almanaque Z/Mexican Cave Fauna in December 15, 2024 (SEE).
[19] Angarita-Sierra (CCE/2018) + Angarita-Sierra et al (Chapter Book, 2019) + Moreno-González, J.A. et al. (American Museum Novitates, 2023) for Colombia.
[20] Chatelliers et al. (Freshw. Biol., 2009), for Clitellata.
[21] Turbanov, I. et al. (Entomological Review, 2016), for former Sovietic Union.
[22] Peter Beron (Historia Naturalis Bulgarica, 2015), for SE Asia to New Guinea, inc. S China and S Japan.
[23] Nicholas, B.G. (The American Midland Naturalist, 1962), for Guatemala to Panama, and Caribbean except Cuba and Jamaica.
[24] Stewart B Peck (Canadian Journal of Zoology, 1999), for Jamaica.
[25] Culver et al. (Conservation Biology, 2021), for U.S.A. and Canada.
[26] Martínez, A. et al (BOOK, 2016), for anchialine caves in Canary Is.
[27] Perez-Moreno, JL et al. (International Journal of Speleology, 2016).
[28] Golovatch, S. et al. (Invertzol, 2018), for European Russia.
[29] Sket et al. (Balkan Biodiversity, 2004), for Balkans.
[30] Zagmajster, M. et al (Global Ecology and Biogeography, 2014), for European Pancrustacea.
White & Culver (2005, 3th edition) is the most complete work on caves in a single publication; however, due to the fact that it is a book written with several contributions, many data are still out of date and others are conflicting with others from the same work. So, whenever necessary, we will use this reference, complemented by others.
5 WORLD DIVERSITY AND NUMBERS
How Many Species of Troglobites [troglobics] Are There? This is the title of an article by Culver & Holsinger (National Speleological Society, 1992), where they estimate there are between 50,000-100,000 troglobic species on the planet. Hobart M. King in Geology (SEE) cites a total of 7,700 spp.
In this work, we have extreme consistent data for Africa, Iran, SE Asia to New Guinea and Vanuatu, China, Canada/U.S.A., Mexico, Guatemala to Panama, Caribbean except Cuba and Jamaica, Cuba, Jamaica, Venezuela, Colombia, Ecuador, Peru, Bolivia and Brazil, and few consistent data for Europe and Asiatic Russia, Central Asia, Middle East except Iran, India Subcontinent, Japan, Australia, smaller Pacific islands except Vanuatu, New Zealand, Guianas and Cono Sur.
In our survey, severely limited due to a huge of problems, lacks of information and inconsistences, we list 10 phylla with true troglobic known for science. Although there is no consolidated data, our work believes that 10,168 troglobics documented in world: approximately 5,000 troglobics from Europe to Asiatic Russia [2, pg. 136], 82 in Middle East to Central Asia and Indian Subcontinent, 1,087 in E & SE Asia to New Guinea, 369 from Africa, 2,858 in New World, and 778 in Australia and Pacific Islands. If we estimate minimally realistically the estimative of M. T. Guzik et al. (2010), with c. 4,140 troglobic spp. from Australia, would be, worldwide, an estimate of 13,390 spp., only 4% in Brazil.
For publications about on extremely diverse caves around the world, see Diversity: Special Issue 'Hotspots of Subterranean Biodiversity'.
We present below numbers from various regions and countries around the world. However, it is important to highlight that these numbers are subject to various biases: many are poorly accurate, do not follow strong criteria regarding the concept of troglobics adopted, some are heavily outdated, much information is unavailable or not compiled, and several conflicts between these numbers can be pointed out. We use them here as reference points for compilation within the most reliable data available.
A hotspots of subterranean biodiversity is a site with 25 or more troglobic, excluding deep soil sites or hyporheic sites unconnected with caves. There are 28 hotspots, that represent almost all subterranean biodiversity hotspots documented thus far at the world scale, in U.S.A. (5), Mexico (1), Brazil (3), Bermuda (1), Spain (3), France (4), Slovenia (2), Croatia (1), Bosnia & Hercegovina (1), Romania (1), China (1), Thailand (1), Vietnam (1), Indonesia (1) and Australia (2). Among these, only Brazil, Australia and Spain has non karstic ones (Culver, Deharveng, Bedos & Pipan, Diversity, 2021).
If we consider only the systems with both troglobites and stygobites, they are reduced to 20: U.S.A. (3), Brazil (3), Spain (1), France (3), Slovenia (2), Croatia (1), Bosnia & Hercegovina (1), Romania (1), China (1), Thailand (1), Indonesia (1) and Australia (1).
If we consider only the systems with 8 or more troglobics in both groups (hyperhotspots), they are reduced to 13: U.S.A. (3), Brazil (1), Spain (1), France (2), Slovenia (2), Croatia (1), Bosnia & Hercegovina (1), Romania (1) and Indonesia (1).
In Brazil, Pedro Cassiano Cave (SEE) and Toca do Gonçalo Cave (SEE), both in Bahia state, are two cave systems with 20 to 25 troglobics.
At least 4 species occur in more than one area: Namanereis cavernicola Solís-Weiss & Espinasa, 1991 in the Caribbean and Mexico, Erpobdella absoloni Johansson, 1913 in Western Europe and the former USSR region (Georgia), Halosbaena acanthura Stock, 1976 in the Caribbean and Venezuela, and Prietella phreatophila Carranza, 1954 in the USA and Mexico.
Europe except former USSR includes among troglobics, 89 Clitellata (39 Tubificida, 29 Lumbriculidae, 10 Enchytraeidae, 5 Haplotaxidae, one Parvidrilidae), 5 Hirudina (one shared with former USSR in Georgia), 2 Namanereis (in Canary Is.).
Europe no has unified checklist of troglobics available. 5,000 troglobics from Europe [2, pg. 136]. Croatia includes 299 troglobionts and 170 stygobitic taxa, including more 100 beetles (Gottstein, S. et al., Natura Croatica, 2002).
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 U.S.A. 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).
Among Pan-Crustacea, Europa includes stygobitic in Ostracoda (114), Copepoda (547), Branchiopoda (7, Cladocera) and Malacostraca (900, being Isopoda 337, Amphipoda 438, Bathynellacea 106, Thermosbaenacea 2, Mysida 3, Decapoda, 16), these by Zagmajster, M. et al (Global Ecology and Biogeography, 2014), and additional Remipedia (2, Canary, Is.). No data for remaining Crustacea.
So far, 99 spp. of troglobic Tricladida [21], members of Annelida/Phyllodocida (2, Canary Is.), Clitellata (92, see topic below), and Actinopteri (2), in Germany and Croatia, one each [14] are known in Europe.
Includes also many troglobics in Gastropoda, Arachnida, Chilopoda, Diplopoda and Pancrustacea. Troglobics Onychophora, Symphyla, Pauropoda are fully unknown in Europe in any form.
[7] lists the troglofauna of Africa, with 396 spp., excluding Actinopteri, but presents some inconsistencies in its numbers. Although it mentions six phyla, only five are listed (page 107). The sum of the clades in Arthropoda totals 344 spp., not 347 as indicated. Three Platyhelminthes are cited, but the list includes six (three Trepaxonemata and three Turbellaria). The sum of Harpacticoida and Cyclopoida results in 59, exceeding the 50 species of Copepoda mentioned.
In the graphs on page 108, three Tricladida and three Neoophora are cited. For the Platyhelminthes, we assume that all references refer to three specific species, and the ambiguous repetitions stem from the repeated inclusion of a group at different hierarchical levels. Chilopoda (2) appears as 'Chilopoda' (1), Scolopendromorpha (1), and Lithobiomorpha (1) in the graphs, but here we adopt Scolopendromorpha (1) and Lithobiomorpha (1) to maintain the numbers as per the original list. 'Blattaria' (4) and 'Blattodea' (1) are shown separately in the graphs, but here we treat them as Blattodea (5). The category Pseudoscorpiones is repeated twice, totaling six species. Amphipoda is mentioned three times, including Bogidellidae, with a total of 72, but we adopt the 69 species initially listed. The graphs indicate 36 Harpacticoidea, 1 Crangonyctoidea, 1 'Copepoda', 1 Calanoidea, and 23 Cyclopoidae, with an actual total of 62; this number is accepted, excluding 1 Copepoda due to lack of details. The mention of 'Malacostraca' (2) is rejected due to insufficient information. The reference to 'Myriapoda' (1) is also rejected, as well as the mention of 'Crustacea' (3) and 'Maxillopoda' (14) due to the lack of details. Finally, the graphs indicate that 'Entognatha' (2) refers to Diplura (2).
Based on these considerations, the direct filtering of [7] results in 357 troglobites in Africa, distributed among the following groups: Platyhelminthes (3), Annelida (2, Clitellata), Mollusca (16), Onychophora (1, in South Africa), Arachnida (32, Trombidiformes 1, Schizomida 1, Pseudoscorpiones 6, Amblypigy 1, Opiliones 2, Araneae 21), Diplopoda (5, 1 Spirostreptida, 1 Chordeumatida, 3 Polydesmida), Chilopoda (2), Ostracoda (4, Podocopida), Copepoda (61, Harpacticoida 36, Cyclopoida 23, Calanoidea 1, Crangonyctoidea 1), Malacostraca (188, Isopoda 71, Amphipoda 69, Bathynellacea 27, Decapoda 15, Thermosbaenacea 5, Spelaeogriphacea 1), Diplura (2), Collembola (5, Poduromorpha 2, Entomobryomorpha 3) and Insecta (36, Coleoptera 23, Diptera 3, Blattaria 5, Orthoptera 1, Dermaptera 1, Hemiptera 1, Hymenoptera 1, Lepidoptera 1).
Also by [7], 32 african countries has troglobics, mainly in Morocco (92), Madagascar (64), Algeria (45), South Africa (34) and Namíbia (16). 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 Moutaouakil, S. et al (Subterranean Biology, 2024).
Addionally, five countries in Africa has 11 troglobic Actinopteri: Cameroon (1:1/1), Namibia (1:1/1), Somalia (2:3/3), D.R. Congo (1:1/1) and Madagascar (2:2/5), by [14], Sendra et al. (Zoological Journal of Linnean Society, 2021) cites 3 Diplura (one more), and [20] cites one more Haplotaxidae from Guinea.
Thus, Africa has 370 troglobic species.
[21] lists 269 spp. from former USSR except Baltic countries (EX-USSR/SS), in Russia (150, Tricladida 3, Annelida 2, Gastropoda 8, Bivalvia 1, Opiliona 3, Pseudoscorpiona 3, Acari 5, Chilopoda 1, Diplopoda 6, Ostracoda 4, Copepoda 21, Bathynellacea 6, Amphipoda 22, Isopoda 19, Decapoda 1, Collembola 21, Diplura 6, Coleoptera 18, mainly in Crimea and Caucasus), Abkhazia (64, Tricladida 1, Annelida 4, Gastropoda 10, Bivalvia 2, Opiliona 3, Palpigrada 1, Chilopoda 1, Diplopoda 4, Copepoda 5, Amphipoda 12, Isopoda 3, Decapoda 4, Collembola 6, Coleoptera 10), Georgia (34, Annelida 3, Gastropoda 3, Bivalvia 1, Opiliona 1, Pseudoscorpiona 2, Diplopoda 1, Ostracoda 1, Copepoda 7, 1 Bathynellacea, Amphipoda 2, Isopoda 3, Decapoda 2, Collembola 2, Diplura 1, Coleoptera 4), Ukraine (11, Bathynellacea 3, Amphipoda 1, Collembola 4, Coleoptera 3, all from Carpathian region), Armenia (1, Bivalvia), Azerbaijan (3, Amphipoda 1, Collembola 2), Kazakhstan (2, Amphipoda), Uzbekhistan (6, Annelida, Gastropoda, Copepoda, Bathynellacea, Isopoda and Collembola one each), Kyrgystan (5, Bathynellacea 4, Isopoda 1), and Turkmenisthan (4, Isopoda 2, Gastropoda and Coleoptera one each). 5 of these species are shared (1 Russia/Abkhazia, 1 Russia/Ukraine, 1 Azerbaijan/Georgia, 2 in Russia/Georgia/Abkhazia). Belarus, Moldova and Tajikistan does no has troglobics.
Troglobitic Clitellata in [20] are absent in [21] and are here counted as part of Western Europe.
Additionally, two more troglobic are known in Turkmenistan: Turkmenocampa mirabilis Sendra & Stoev (Diplura), found in Kaptarhana cave, in Lebap province (Sendra, A. et al., Subterranean Biology, 2017), and Paracobitis starostini Parin, 1983 (Actinopteri, Nemacheilidae), by [14].
Thus, troglobics in EX-USSR/SS are 276, being Tricladida (4), Annelida (10, one shared with W Europa), Gastropoda (22), Bivalvia (4), Acari (5), Pseudoscorpiona (5), Opiliona (6), Palpigrada (1), Chilopoda (2), Diplopoda (11), Ostracoda (5), Copepoda (30), Bathynellacea (15), Amphipoda (40), Decapoda (7), Isopoda (29), Diplura (8), Collembola (35), Coleoptera (36) and Actinopteri (1).
Middle East and Afganisthan no has unified checklist of troglobics available. At least 12 troglobics, one Diplura (Sendra et al., Zoological Journal of Linnean Society, 2021, in Lebanon), three Namanereis (Annelida/Nereididae), one in Oman and two in Socotra [2], two Clitellata in Oman [20], and six Actinopteri, in Iraq (2:3/3) and Oman (1:1/3), by [14].
Iran includes 51 troglobics by [8], being 46 aquatic and five terrestrial. However, we were only able to identify 48 of them in this post: 24 Amphipoda, 12 Copepoda, 3 Isopoda [8], Araneae and Diplopoda one each [9], one Clitellata (Erpobdella borisi Cichocka & Bielecki, 2015), one Gastropoda and (2:2/)5 Actinopteri [14].
Indian Subcontinent (Pakistan, India, Bangladesh, Nepal, Butan and Maldivas) no has unified checklist of troglobics available. 21 troglobic Actinopteri are known from this regions, in India (7:10/21), by [14], and one Diplura, also in India (Sendra et al., Zoological Journal of Linnean Society, 2021).
Nippon no has unified checklist of troglobics available. Nearly 200 spp. of troglobics are known from the groundwaters of Nippon. Most of these troglobic species, 16 of 77 genera, and what is more, 4 of 47 families are endemic to Nippon. Uchidastygacaridae, Nipponacaridae, and Kantacaridae are endemic acaridan families of Nippon (Matsumoto, Kôichi, International Journal of Speleology, 1976). Two Clitellata [20] and one Diplura (Sendra et al., Zoological Journal of Linnean Society, 2021) are troglobics in Nippon. (1:1/)5 troglobic Actinoperi, all in Luciogobius (Gobiidae), are known from Nippon [14].
We considere here 710 troglobics in China, in Gastropoda (2), Araneae (104), Pseusocorpiona (40), Copepoda (23), Bathynellacea (1), Amphipoda (26), Decapoda (23), Isopoda (18), Chilopoda (1), Diplopoda (114), Collembola (42), Orthoptera (40), Coleoptera (159), Hemiptera (1), Diptera (2), these by [2], Annelida (1, see notes in Annelida description, Hirudina. [2] cites two species, however none details among the secodn was found in literature), Diplura (5) by Sendra et al. (Zoological Journal of Linnean Society, 2021), and Actinopteri (107 in three families), by [14], near a half are fishes, beetles and spiders
[2] cites one cave-dwelling Anura in China, however is fully rejected here as troglobic, being excludes in this count.
[22] lists all the non-Actinopteri troglobionts from Southeast Asia to New Guinea. Their list contains inconsistencies between the manual counts and the numbers provided in the abstract: 16 fewer species are counted in Southeast Asia and 6 fewer in New Guinea. Considering only the 11 politically recognized Southeast Asian countries (Myanmar, Thailand, Laos, Vietnam, Cambodia, Malaysia, Cingapore, Indonesia, Philippines, Brunei, East Timor, and Papua New Guinea), excluding China and Japan members, we total 146 troglobic species, being Phyllodocida (1, Namanereis from Ppaua New Guinea), Gastropoda (1), Araneae (13), Pseusoscorpiones (4), Scorpiona (2), Thelyphonida (1), Opiliones (3), Acari (4), Diplopoda (15), Amphipoda (3), Isopoda (22), Decapoda (22), Diplura (2), Collembola (23), Orthoptera (1), Blattaria (8), Coleoptera (18), Hymenoptera (1) and Diptera (2).
Additionally, two troglobics Diplura are cited for this region, in Myanmar (1) and Papua New Guinea (1), in Sendra et al. (Zoological Journal of Linnean Society, 2021), and 29 troglobic Actinopteri are known from this regions, in Myanmar (1:1/1), Thailand (4:5/9), Laos (2:3/3), Vietnan (3:6/7), Malaysia (1:1/1), Philippines (2:2/3), Indonesia (3:3/3) and Papua New Guinea (1:1/2), by [14].
By country/or region: Myanmar (4), Thailand (47), Laos (17), Cambodia (1), Vietnan (24), Philippines (14), continental Malaysia (4), Malaysian Sarawak (18), Malaysian Sabah (1), Sumatra and adjacent Batu (6), Java and adjacent Sumba (5), Indonesian Kalimatan (4), Sulawesi (7), Moluccas (1) and New Guinea (25).
Thus, SE Asia includes 177 troglobic species.
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. Towakkalak cave in Sulawesi has 36 spp. of troglobics (SEE), but non counted here.
Australia no has unified checklist of troglobics available, and the lack of precision in the numbers from Australia is absurdly complicating and confusing. [2] cites troglobic Amphipoda (144), Palpigrada (1) and Scopiona (1). We combined these numbers with the following numbers from [12]: Aracnida (101, for uniquely didactic effect, all assignated for Araneae), Myriapoda (19, for uniquely didactic effect, all assignated for Diplopoda), Hexapoda (164, for uniquely didactic effect, all assignated for Coleoptera), 'terrestrial Crustacea' (45, for uniquely didactic effect, all assignated for Isopoda), Gastropoda (4), Ostracoda (74), Copepoda (83), Bathynellaceae (83) and Isopoda (49), who discussed the troglofauna of Western Australia. We excluded from this second work the imprecise placements or defased numers for some groups: other Stygofauna' (30).
Additionally for troglobics, there are Hanseniella magna Scheller in Tasmania (Symphyla, SEE), one more Diplura (Sendra et al., Zoological Journal of Linnean Society, 2021), one Clitellata [20], Namanereis pilbarensis Glasby, Fiege & Van Damme, 2014 (Nereididae) in Pilbara region in Australia (SEE), (2:2/)3 Actinopteri (in Milyeringidae and Synbranchidae) in N Australia [14], Kumonga (1, Remipede) from NW Australia, unique in this class in southern Hemisphere, Halosbaena tulki Poore & Humphreys, 1992 is Western Australia (Wikipedia) and certainly 3 spp. Anaspidacea (Wikipedia).
In these fragmented and imprecise numbers, we estimate the diversity of troglobites in Australia at 780 spp.
Pacific region except Papua New Guinea and New Zealand no has unified checklist of troglobics available. At least one species of troglobic Nemertea is known in New Zealand, Exosphaeroides quirosi Jaume et Queinnec, 2007 (Sphaeromatidae, Isopoda) for Vanuatu [22], Phreodrilus subterraneus Beddard, 1891 (Clitellate/Phreodrilidae) also in New Zealand [20], one Namanereis (Annelida/Nereididae) in New Zealand and Fiji [2], ona Gastropoda in New Zealand [2, Hydrophrea], one Actinopteri from Guam (1:1/1), by [14], and two Anaspidacea in New Zealand (Stygocarella pleotelson Schminke, 1980 and Stygocaris townsendi Morimoto, 1977, SEE).
By [2], number of described non-Actinopteri troglobics from the U.S.A. and Canada, which includes Alaska and Hawaii, is (124:288/)1,343 spp., representing 5 phyla, 15 classes and 51 orders (1/2 are spiders, diplopods, pseudoscorpiones or beetles, and 1/5 amphipodes or isopods). Additionally, one uncommon Annelida (Parvidrilus strayeri Erséus, SEE), 5 more Diplura (and one more genera, 1 in Canada, 4 in U.S.A., Sendra et al., Zoological Journal of Linnean Society, 2021) and 22 spp. of Actinoperi is known from U.S.A. [14]. Thus, U.S.A. (including Hawaii and Alaska) and Canada has 1,371 troglobics.
In California there are 102 troglobionts (terrestrial cave obligates), 12 stygobitics (aquatic cave obligates), and 32 phreatobites (obligate groundwater forms), by Elliot, R.W. et al (Proceedings of the California Academy of Sciences, 2019). Missouri has 82 troglobics (67 described, 15 undescribed), including 49 aquatic and 33 terrestrial species. The aquatics include 30 described and six undescribed stygobitics, plus 13 described phreatobites. The terrestrials include 24 described and nine undescribed species. Six of the troglobics (four described) may actually be troglophiles, edaphobites or neotroglobics (Elliot, R.W., Journal of Cave and Karst Studies, 2007).
MEXICO (538)BRAZIL/U.S.A. COMPARATION
Neither Brazil nor the U.S.A. have troglobics in Nematoda and Cnidaria.
Troglobic lineages in Brazil unknown as troglobics in U.S.A. are Porifera, Nemertea, Onychophora, Bivalvia, Pauropoda, Symphyla, Blattodea and Hymenoptera.
Troglobic lineages in U.S.A. unknown as troglobics in Brazil are Platyhelminthes/Alloeocoela (1/1), Thermosbaenacea (1/1), Schizomida (2/2) and Amphibia (1:2/11).
At commom groups, Brazil and U.S.A. are fully tied in Scorpiona, Zygentoma, Dermaptera and Diptera.
At commom groups, U.S.A. surpasses the Brazil in the following groups: Tricladida (3G, 18s), Annelida (1F, 4G, 12s), Gastropoda (7G, 10s), Acari (2F, 9G, 18s), Pseudoscorpiona (4F, 23G, 109s), Araneae (51s), Opiliona (1G, 8s), Diplopoda (13G, 67s), Oligostraca/Ostracoda (1F, 10G, 15s), Copepoda (2F, 5G, 13s), Decapoda (2F, 6G, 36s), Bathynellacea (1F, 7G, 18s), Amphipoda (5F, 14G, 161s), Diplura (2G, 5s), Collembola (9s), Coleoptera (5F, 22G, 221s).
At commom groups, Brazil surpasses the U.S.A. in the following groups at all levels: Chilopoda, Orthoptera, Hemiptera and Actinopteri.
Mexico has 538 spp. of troglobics in the following groups, by [18]: Porifera (1, Legendre et al., Subterranean Biology, 2023, undescribed), Platyhelminthes (2:2/6, 4 Continenticola, 2 Cavernicola), Annelida (3/5, one Errantia, two Oligochaeta, two Branchibdellida), Gastropoda (3:5/9), Remipedia (3, Xibalbanus), Copepoda (5:8/12), Ostracoda (4, three genera of three families in two orders), Amphipoda (2:22 spp.), Thermosbaenacea (1, Tulumella unidens from Quintana Roo), Mysida (3:3/7), Decapoda (27 spp.: 17 shrimps in six families, 5 crayfishes at Cambaridae, and 5 crabs in two families), Isopoda (11/52), Collembola (9:/42), Diplura (8, two Litocampa, two Juxtlacampa, one Oncinocampa and three Tachycampa, Sendra et al., Zoological Journal of Linnean Society, 2021), Zygentoma (3/13), Orthoptera (6, all Phalangospsidae), Coleoptera (44 spp., Carabidae/30, Dytiscidae/1, Histeridae/4, Leiodidae/8, and Ptinidae/1), Hemiptera (3, all Fulguromorpha), Diplopoda (15:/73, at six orders), Chilopoda (5, two Lithobiidae, 3 in Newportia), Trombidiformes (2), Pseudoscorpiona (40 spp. in 5 families), Araneae (11:59 spp.), Opiliona (4:16 spp.), Amblypygi (11, all Phrynidae), Schizomida (23, 14 in Hubbardiidae, 9 in Protoschizomidae), Ricinulei (11, all in Pseudocellus), Scorpionida (4:8/13), and Actinopterigii (5:6/20 spp. at 5 orders).
Mexico no has described troglobic in Nematoda, Nemertea, Onychophora, Mollusca/Bivalvia, Bathynellacea, Spelaeogriphacea, Blattodea, Dermaptera, Diptera, Hymenoptera, Pauropoda, Symphyla, Palpigradi or Amphibia. Brazil has more species than Mexico in all groups except in Remipedia, Copepoda, Ostracoda, Amphipoda, Thermosbaena, Mysida, Decapoda, Zygentoma, Coleoptera, Diplopoda, Pseudoscorpionida, Araneae, Amblypygi, Schizomida, Ricinulei and Scorpionida, all in Arthropoda.
The only consistent reference on the troglobitic 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.
By [23], only eight troglobics occur from Guatemala to Panama, namely: Otostigmus cooperi Chamberlin, 1942 (Chilopoda, Panama), Pselliodes harveyi Chamberlin, 1942 (Chilopoda, Panama), Pselliphora cavincola Chamberlin, 1918 (Chilopoda, Trinidad), Calymmodesmus inquinatus Causey, 1960 (Diplopoda, Guatemala), Lepidocyrtus usitatus Folsom, 1927 (Collembola, Panama), Ptomaphagus giaquintoi Jeannel, 1936 (Coleoptera, Alta Verapaz, Guatemala), Trombicula cavernarum Ewing,, 1933 (Trombidiformes, Panama) and T. trifurca Ewing, 1933 (Trombidiformes, Panama). Additonally, one Diplura in Guatemala (Sendra et al., Zoological Journal of Linnean Society, 2021), one troglobitic Actinopteri is known from Belize, Rhamdia typhla Greenfield, Greenfield, and Woods, 1982, in Heptapteridae [14], and one Remipede occur in Belize (Xibalbanus cokei Yager, 2013, SEE).
Thus, in America Central are known 11 troglobitics. Deleva, S. et al. (Subterranean Biology, 2023) discusses the cavernicolous fauna of Costa Rica and confirms that there are no troglobitic 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.
[13] cites 112 troglobics in Cuba, all Arthropoda: Acari (12), Scorpiones (1), Pseudoscorpiones (3), Amblypygi (1), Schizomida (5), Ricinulei (4), Opiliones (4), Araneae (7), Chilopoda (2), Remipedia (1), Ostracoda (5), Copepoda (18), Mysidacea (5), Isopoda (7), Decapoda (13), Amphipoda (3), Zygentoma (3), Collembola (7), Orthoptera (4), Blattoidea (1) and Coleoptera (6). Additionally, six Thermosbaenacea (Wikipedia), 5 Clitellata [20], two Chilopoda (see topic about Chilopoda) and nine Lucifuga (Bythitidae, Ophidiiformes, Actinopteri) are reported [14].
Thus, Cuba has 134 troglobic species in our survey.
Troglobic in Jamaica contains 43 spp. troglobics, being 27 of troglobionts in Onychophora (Speleoperipatus spelaeus Peck, 1975), Araneae (9), Opiliona (2), Pseudoscorpionida (4), Schizomida (1), Isopoda (1), Collembola (2), Blattodea (1), Hemiptera (2) and Coleoptera (4), and 16 stygobitics, in Tricladida (1), Ostracoda (3), Mysida (2), Thermosbaenacea (1), Decapoda (2), Amphipoda (4) and Isopoda (3), by [24].
GUIANAS (0)The only consistent reference on the troglobitic 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 troglobics for this area, namely: Porcellionides minutissimus Van Name, 1936 (Isopoda, Bahamas), 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), Maymena bruneti Gertsch, 1960 (Araneae, Trinidad), Stygiomysis holthuisi Gordon, 1960 (St. Marteen, Mysida), Calycuoniscus spinosus Collinge, 1917 (Isopoda, Trinidad, West Indies) and Philoscia bermudensis Dahl, 1892 (Isopoda, Walsingham Cave, Bermuda).
Additionally, 39 more troglobics are confirmed for this area: 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], 17 Remipedia in Bahamas (7/13), Turks y Caicos (4/4) and Republic Dominicana (1/1), 10 Thermosbaenacea widely in many islans in region (9 exclusives and one also in Venezuela, Wikipedia), three Actinopteri, Rhamdia urichi Mees, 1974 (Heptapteridae), from Trinidad and Tobago, and two Lucifuga (Bythitidae) from Bahamas [14].
Thus, in West Indies except Jamaica and Cuba are known 48 troglobitics.
VENEZUELA (47)[11] cites possibly Lepidocampa (Diplura), Eukoenenia (Palpigrada) and Modisimus (Araneae) as troglobics in French Guiana, however their troglobic status in unknown, and all are rejected here.
COLOMBIA (38)The most recent listing for Venezuelan troglofauna, [17], dates back to 1 and a half decades. There are 46 troglobics listed for the country: nine Acari (4 families, none with troglobics in Brazil), four Opiliona (all in Trinella, Agoristenidae), two Amblypygi (both Charinus), one Scorpionida (Taurepania trezzii, Chactidae), one Diplopoda (Trichopolydesmidae), one Collembola (Onychiuridae), one Copepoda/Cyclopidae, two Ostracoda (both Cyprididae), one Bathynellaceae (Parabathynellidae), six Isopoda (in six families), three Amphipoda (in three families), two Decapoda (both crabs Pseudothelphusidae), two Orthoptera (both Rhaphidiophoridae), one Blattodea (Paranocticolla venezuelana), three Coleoptera (in three families), and seven fishes (in 4 families). Cyathura univam (Isopoda) and Metaniphargus venezuelanus (Amphipoda) are anchialines.
Additionaly, [20] cites more one species for Venezuela, Krenedrilus papillatus Dumnicka, 1983 (Annelida/Tubificidae), there are one Thermosbaenacea (Halosbaena acanthura Stock, 1976, Wikipedia), and [14] cites only five fishes, data accepted here. Thus, 47 troglobics occur in Venezuela.
ECUADOR (7)[19] cites 62 troglobics in Colombia, in Nematoda (1, Rhabditida), Annelida (2, Haplotaxide, Enchytraeidae one each), Tityus grottoedensis (Scorpionida), Opiliona ((7:)7), Amblypygi (2, Phrynidae and Paracharontidae), Araneae (16, in 11 families, Barychelidae, Araneidae, Anyphaenidae, Lycosidae, Tetragnathidae, Pisauridae without Brazilian troglobics), Chilopoda (3, undescribed Scolocryptops and 2 Psilliodes), Diplopoda (3), Isopoda (1, Armadillidium sp.), 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, SEE), 4 Opiliona (Angarita-Sierra et al, Chapter Book, 2019), one Ephemeroptera (Angarita-Sierra et al, Chapter Book, 2019), Cavernicola pilosa (SEE) and Strudvelia cinctipes (SEE). Additionally, [14] recognizes 10 troglobic Actinopteri in country.
With these exceptions, Colombia has, with some consistency, 38 spp. of troglobics: Nematoda (1, Rhabditida), Opiliona (3), Amblypygi (2, Phrynidae and Paracharontidae), Araneae (3), Chilopoda (1, undescribed Scolocryptops), Diplopoda (3), Isopoda (1, Armadillidium sp.), 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.
PERU (3)Ecuador no has unified checklist of troglobics available. Consistently, ony 7 troglobics ocur in Ecuador: Marionina ecuadoriensis Righi and Hemienchytraeus mauriliae Righi (Clitellata/Enchytraeidae) by [20], Tayos ashmolei Reddell & Cokendolpher, 1984 (Schizomida, SEE), Troglotayosicus vachoni Lourenço, 1981 (Scorpiona/Troglotayosicidae), Ischnoptera peckorum Roth, 1988 (Ectobiidae, Blattaria) and two Actinopteri, Astroblepus pholeter Collette 1962 and one Chaetostoma undescribed species [14].
BOLIVIA (8)Only three spp. in Peru the following can be considered troglobics: Acheroxenylla lipsae Palacios-Vargas, 2020 (Collembola, Poduromorpha, Hypogastruridae), Astroblepus riberae Cardona and Guerao, 1994 (Actinopterygii, Siluriformes, Astroblepidae) and Caecopactes chullachaqui Campos-Filho, Sfenthourakis and Bichuette, 2024 (Isopoda, Scleropactidae) are known [16].
ARGENTINA, PARAGUAY, URUGUAY AND CHILE (1)Bolivia no has unified checklist of troglobics available. Six troglobics 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 [14] are troglobic in Bolivia, totalling 8 troglobitics in country.
Without a unified checklist of troglobics available, our survey counted 8 troglobics for the region: six Anaspidacea (SEE | SEE), one Hemiptera (Notolathrus sensitivus Remes Lenicov, 1992, SEE) from Argentina, and one Actinopteri, Silvinichthys bortayro Fernandez and De Pinna 2005 (Trichomycteridae), known for from Argentina [14].
A TABLE OF ALL NUMBERS USED IN THIS POST, UPDATES IN DECEMBER 26, 2024
6 BRAZILIAN NUMBERS
This list of cave-obligate taxa is in a continual state of flux, as new species are added through new species descriptions, some are removed by being synonymized with other species, and others are reclassified as troglophiles or stygophiles when records from surface habitats or other new evidence is obtained. Moreover, we recognize that the inclusion or exclusion of particular taxa may be contentious for some biospeleologists. 678 spp. (all Brazilian endemics except Phreatobius sanguijuela), being:
∷ 338 describeds, being:
▪ 325 state endemics.
▪ 13 in two or more states.
SOURCE
These 338 spp. describeds includes:
142 describeds in [1].
[1] cites 150 spp., but 8 are rejected here as troglobits: Troglorhopalurus lacrau (BA, Scorpionida), Isoctenus corymbus (Ctenidae), Lygromma ybyguara (Prodidomidae), and Leonardossia hassalli (PA, Isopoda), rejected by [6]; Metagonia potiguar (RN, Araneae), non cited at Bento et al. (Biodiversity and Conservation, 2022); Ochyrocera ibitipoca (MG, Araneae), rejecyed in A. Brescovit & I. Cizauskas (Arachnology, 2018); and Endecous apterus and E. peruassuensis (rejected in Carvalho, P.H.M. et al, Zootaxa, 2023).Discocyrtus pedrosoi (BA, Opiliona), rejected in [6], is accepted here.
196 describeds latter [1], or after but non cited in world [A].
∷ 340 undescribeds [B], being:
▪ 339 state endemics.
▪ one shared for two states (a Cirolanidae in Rio Grande do Norte and Ceará).
NOTES AND OBSERVATIONS
OTHER REJECTED SPECIES (1): Pseudonannolene canastra (Diplopoda), synonimized under P. ambuatinga (Iniesta, Bouzan & Brescovit, European Journal of Taxonomy, 2023).
TROGLOBICS AWAITING CARD ANALYSIS (0).
SPECIES COMPLEX BY UNCOMMON HUGE DISTRIBUTION AMONG TROGLOBICS (2, possibly 5): Pseudochthonius biseriatus, described for N Minas Gerais state, also collected in Catimbau National Park, Pernambuco state (SEE), possibly two species; and Pseudochthonius strinatii, known from São Paulo, Paraná and Minas Gerais states, possibly three species (SEE).
SPECIES WITHOUT TROGLOBIC NON CONFIRMED (2): Whiteheadiana thaisae (Carabidae, Benumea et al., Zootaxa, 2023) and Pseudochthonius itakuatiara (Pseudoscorpiona, Zoological Studies, 2024).
QUESTIONS: would Kadiweuoniscus rebellis be the species referred to as Oniscoidea 1, in Cordeiro et al. (Biota Neotropica, 2014)? No indication denying or affirming this was found in our research, thus here we consider them as distinct taxa.
[A] 196 formal described species not mentioned in [1], almost all described later:
PORIFERA (1)
PORIFERA (1): Arinosaster patriciae (MT, Neotropical Biology and Conservation, 2021).
PLATYHELMINTHES (6)
PLATYHELMINTHES (8): Girardia arenicola, G. corumbataiensis, G. nobresis, G. paucipunctata, G. spelaea (SP, MS, BA, Zoologischer Anzeiger, 2021), Girardia patiensis, G. alba (BA, Zoologischer Anzeiger, 2025) and Sluysia triapertura (RN, Invertebrates Systematics, 2018).
MOLLUSCA (5)
BIVALVIA (1): Eupera troglobia (TO, Subterranean Biology, 2022).
GASTROPODA (6): Habeastrum parafusum, H. omphalium (MS, Zootaxa, 2019), Gastrocopta sharae (GO, Zoosystematic Evolution, 2017), Spiripockia umbraticola (BA, Journal of Natural History, 2021), Idiopyrgus eowynae and I. meriadoci (BA, Zoosystematics and Evolution, 2024).
ANTHROPODA/ARACNIDA (68)
TROMBIDIFORMES (1): Leptus sidorchukae (MG, Zootaxa, 2019).
PSEUDOSCORPIONA (19): Spelaeobochicha goliath (MG, Zootaxa, 2018), S. mahnerti (MG, Zootaxa, 2020), Pseudochthonius ramalho, P. olegario, Spelaeochernes altamirae, S. armatus, S. bahiensis, S. dentatus, S. dubius, S. eleonorae, S. gracilipalpu, S. pedroi, S. popeye (BA, MG, SP, GO, MS, MG, BA, SE, PR, SC, Arachnology, 2022 + Revue Suisse de Zooologie, 2001), Pseudochthonius koinopoliteia, P. diamachi, P. pali (Prado & Ferreira, Zootaxa, 2023), Pseudochthonius lubueno (BA, Zoologia, 2023), Pseudochthonius aware (BA, Zoological Studies, 2024) and Pseudochthonius limettioides (MG, Zootaxa, 2024).
PALPIGRADES (12): Allokoenenia canhembora, A. stygia (BA, PA, European Journal of Taxonomy, 2022), Eukoenenia eywa, E. navi, E. neyriti (MG, Invertebrate Systematics, 2018), E. jequitai, E. lundi, E. magna (MG, Zootaxa, 2020), E. audax (GO, Zootaxa, 2020), E. igrejinha (MG, Journal of Arachnology, 2019), E. ibitipoca (MG, Zootaxa, 2019) and E. mocororo (MG, Zootaxa, 2022).
AMBLYPYGI (5): Charinus cearensis, C. diamantinus, C. euclidesi, C. puri and C. renneri (many states, EJT, 2021).
OPILIONA (4): Iandumoema cuca, I. gollum, I. stygi (MG, Invertebrate Systematics, 2020) and Spaeleoleptes gimli (BA, EJT, 2024).
ARANEAE (30): Ctenus igatu (BA, Journal of Aracnology, 2022), Paracymbiomma caecus (PA, Zootaxa, 2018), Paracymbiomma bocaina (PA, Zootaxa, 2018), Paracymbiomma pseudocaesus (PA, Dissertation, 2017), six Matta (MG, Zootaxa, 2019), Carajas paraua, Tisentnops mineiro, T. onix (PA, MG, ZooKeys, 2016), Loxoscelis troglobia (BA, Zootaxa, 2018), Loxosceles bodoquena (MS, EJT, 2024), Loxosceles boqueirao (BA, EJT, 2024), Tonton itabirito, T. matodentro (MG, ZooKeys, 2013), Ochyrocera dorinha, O. rosinha, O. ungoliant (MG, PA, Subterranean Biology , 2021), Ochyrocera ritxoco, O. ritxoo (PA, Zookeys, 2021), Speocera babau, S. pinima, S. piquira (PA, Zootaxa, 2022), Brasilomma enigmatica (MG, Zootaxa, 2012), Indiani gaspar (MG, Zootaxa, 2020), Pinelema elinae (BA, Zootaxa, 2024) and Paleotoca diminas (MG, Taxonomy, 2024).
ANTHROPODA/MILIPEDES (7)
CHILOPODA (2): Schendylops janelao (MG, Zootaxa, 2019), Plutogeophilus jurupariquibaba (SP, Organisms Diversity & Evolution, 2023) and Cryptops didi (SP, Journal of Natural History, 2024).
DIPLOPODA (5): Dobrodesmus mirabilis (BA, Zootaxa, 2016), Phaneromerium troglopterygotum (BA, Zootaxa, 2022), Strongylosomides troglobius (BA, Zootaxa, 2022), Onciurosoma troglobium (BA, Arthropoda Selecta, 2022) and Cayenniola albaserrata (BA, Zootaxa, 2024).
ANTHROPODA/MALACOSTRACA (47)
AMPHIPODA (1) Spelaeogammarus ginae (BA, Nauplius, 2022).
ISOPODA (45): Amazoniscus spica, Alboscia jotajota, Metaprosekia igatuensis (PA, SP, BA, European Journal of Taxonomy, 2020), five Xangoniscus (BA, MG, Zootaxa, 2020), Xangoniscus lapaensis, X. loboi (BA, Subterraneam Biology, 2022), Chaimowiczia tatus, C. uai (BA, MG, Subterranean Biology, 2021), Chaimowiczia obybytyra (BA, Subterranean Biology, 2022), Iansaoniscus leilae, I. paulae (BA, Subterranean Biology, 2022), Pectenoniscus montalvaniensis, P. juveniliensis, P. iuiuensis, P. carinhanhensis, Pectenoniscus santanensis, P. morrensis (BA, MG, Nauplius, 2020), Pectenoniscus lilieae, Benthana xiquinhoi (BA, Zootaxa, 2019), Pectenoniscus fervens, P. pankaru (PI, BA, ZooKeys, 2022), Spelunconiscus septemlacuum (MG, Nauplius, 2022), Cylindroniscus platoi (MG, Zootaxa, 2022), Gabunillo aridicola (CE/RN, Zootaxa, 2010), Benthana alba, Benthanoides amazonicus, B. tarzan (PA, Zootaxa, 2023), Pectenoniscus monsviridis, P. revelatus, P. archaeos, P. sepultus, P. caesarius (BA, Studies on Neotropical Fauna and Environment, 2023), Caraiboscia jabutiensis, Gabunillo enfurnado, Venezillo moreirai, V. limai (many states, Zoosystema, 2023), Kadiweuoniscus rebellis (MS, ZooKeys, 2024), Circoniscus mendesi, C. xikrin (PA, EJT, 2024), Ctenorillo iuiuensis (BA, Zoosystema, 2024), Trichorhina baiana (BA, Biota Neotropica, 2023), Xangoniscus paiabare, X. tymaopeba, X. ykanhema, X. puku (BA, Tropical Zoology, 2024), Xangoniscus antiquus, X. chaimowiczi and X. jonasi (BA, Subterranean Biology, 2024).
DECAPODA (1): Aegla charon (SP, Nauplius, 2017).
ANTHROPODA/COLLEMBOLA (24)
COLLEMBOLA (24): Arrhopalites glabrofasciatus, Pseudosinella guanhaensis (MG, Zootaxa, 2018), Pararrhopalites ubiquum (MG, Neotrop. Entomol., 2018), 10 in Trogolaphysa, 9 in Pseudosinella (PA, MG, SP, Insects, 2020, and PRE-PRINT, 2021), Troglobentosminthurus luridus (BA, Insects, 2022) and Pararrhopalites sideroicus (MG, Florida Entomologist, 2014).
ANTHROPODA/INSECTA (22)
ORTHOPTERA (4): Endecous infernalis (BA, Zootaxa, 2023), Erebonyx catacumbae (BA, Zootaxa, 2021), Endecous troglobia (MG, Zootaxa, 2020) and Endecous vitreus (MS, Zootaxa, 2023).
COLEOPTERA (11): Ardistomis ferreirai (PA, Zootaxa, 2018), Coarazuphium auleri (MT, Studies on Neotropical Fauna and Environment, 2021), C. bambui (BA, Zootaxa, 2022), C. lundi (MG, Zootaxa, 2020), C. xingu, C. xikrin, C. kayapo (PA, Subterranean Biology, 2022), Metopioxys carajas (PA, Zootaxa, 2019), Oxarthrius aurora, O. inexpectatus (TO, BA, Zootaxa, 2018) and Perigona spelunca (MG, EJT, 2022).
DERMAPTERA (1): Mesodiplatys falcifer (BA, ZooKeys, 2018).
DIPTERA (1): Deanemyia maruaga (AM, Mem. Inst. Oswaldo Cruz, 2011).
HEMIPTERA (5): Ferricixius michaeli, F. goliathi (MG, Zootaxa, 2023), Spelaeometra gruta, Cephalometra pallida (BA, Tijdschrift voor Entomologie, 2018), Spelaeometra hypogea (BA, Animals, 2023).
CRANIATA (1)
ACTINOPTERI (1): Phreatobius sanguijuela (RO/Bolivia, Fauna Cavernicola do Brasil, 2022).
[B] 340 spp. identifiable as troglobits but not yet formally described, but with certainty or very high signs of troglomorphisms, distributed along following state/references below. Codes in bold, e.g. [MG/4], will be used later to cite these works throughout the group discussion.
MGCordisburgo region in MG (35 spp., [MG/1], SEE, Cordisburgo and Curvelo; SUPPLEMENTARY MATERIAL).Arinos in MG (7, [MG/7], SEE).:: 1 of the originally 4 spp. undescribeds in MG/8 have already been published: Pseudochthonius olegario (former Chthoniidae sp. 4, SEE).Matozinhos in MG (4, [MG/9], SEE).Vazante in MG (5, [MG/10], SEE).Bambuí system in MG (unknown, SEE).TOTocantins (6 spp., Dianópolis, [TO/1], Ferreira et al., CECAV, 2016).MSMato Grosso do Sul (31 spp., [MS/1], Cordeiro et al., Biota Neotropica, 2014).RNRio Grande do Norte state (51 spp., [RN/1], Bento et al, Biodiversity and Conservation, 2022; EXCEL LIST); Potiberaba porakuara also in Ceará).ESEspírito Santo state (10 spp., [ES/1], SEE).GOSão Domingos region in GO (10 spp., [GO/1], SEE).BA:: 4 of the originally 22 spp. undescribeds in BA/1 have already been published: Pseudochthonius koinopoliteia (former Chthoniidae sp.1, SEE), Endecous infernalis (former Endecous sp., SEE), Troglobentosminthurus luridus (former Sminthuridae sp.1, SEE) and Cayenniola albaserrata (former Chelodesmidae sp. 1, SEE).:: 1 of the originally 33 spp. undescribeds in BA/2 have already been published: Pectenoniscus iuiuensis (former Styloniscidae sp. 3, SEE).Igatu region in BA (29 spp., [BA/3], SEE).Carinhanha region in BA (10 spp., [BA/5], SEE).
Carinhanha region in BA (Pedro Cassiano cave, 13 spp., [BA/6], Acta Oecologica, 2025).
SPAlto do Ribeira region in SP (six Potamolithus (Lithoglyphidae), [SP/1], Maria Elina Bichuette & Eleonora Trajano, Subterranean Biology, 2018).MIXEDCampo Formoso region in BA (15 spp. in BA, [BA/4], SEE) and Iporanga region in SP (13 spp. in SP, [SP/2]).:: 1 of the originally 14 spp. not described in SP/2 have already been published: Plutogeophilus jurupariquibaba (Chilopoda, SEE).:: 1 of the originally 16 undescribeds in BA/4 have already been published: Erebonix catacumbae (former Phalangospsidae sp. n., SEE).
An important note! The references above cite many more species than those counted here - this surplus is composed almost entirely by species that, at the time of the articles, had not yet been described but that were later described. These species are strictly troglobic species mentioned in the articles and whose formal description our website has not yet been able to confirm. Furthermore, the amount of species not described in each work that this blog was able to collect may not be all the posts in the work, since many of them only mention a very vague amount, and bring images of some; in this case, we extract as accurate as possible from undescribed species.
BY GROUP
Platyhelminthes (4) ‣ SP (1), MS (1), RN (1), MG (1).
Nemertea (2) ‣ SP (1), MG (1).
Annelida (6) ‣ MS (2), MG (2), RN (2).
Mollusca (15) ‣ BA (7), SP (6), MS (2).
Onychophora (1) ‣ MS (1).
Copepoda (4) ‣ RN (2), SP (1), MG (1).
Ostracoda (1) ‣ RN (1).
Amphipoda (5) ‣ RN (4), MS (1)
Bathynellacea (1) ‣ MS (1).
Isopoda (48) ‣ BA (18), MG (13), RN (9, 1 non-endemic), SP (2), ES (2), GO (2), MS (1), TO (1), CE (1, non-endemic).
Collembola (50) ‣ MG (16), BA (13), RN (9), MS (5), SP (4), GO (3).
Diplura (9) ‣ MG (3), RN (3), BA (3).
Insecta (44)
Zygentoma ‣ BA (3), ES (2), MG (1).
Coleoptera ‣ MG (10), BA (8), RN (2), MS (1), SP (1), GO (1), TO (1).
Blattodea ‣ BA (4), MG (2).
Hemiptera ‣ MS (1), BA (3).
Hymenoptera ‣ BA (1), TO (1).
Pauropoda (1) ‣ MG (1).
Symphyla (2) ‣ SP (1), MG (1).
Diplopoda (53) ‣ BA (16), MG (12), SP (11), MS (4), ES (3), RN (3), GO (3), TO (1).
Chilopoda (14) ‣ BA (7), RN (6), SP (1).
Acari (7) ‣ BA (4), MG (2), MS (1).
Araneae (32) ‣ BA (13), MG (7), RN (7), ES (1), SP (1), MS (1), TO (2).
Pseudoscorpionida (15) ‣ BA (6), MG (6), RN (1), SP (1), GO (1).
Palpigrada (7) ‣ BA (5), MG (1), SP (1).
Opiliona (16) ‣ MS (6), MG (4), BA (3), ES (2), RN (1).
Actinopteri (5) ‣ MS (3), BA (2).
Some inventories result in confirmation of the absence of troglobics in some locations, such as Martins (RN, Araujo et al., Espeleo-Tema, 2017). Some works also maybe inconclusive in quantity. Ferreira, Zampulo & Souza-Silva (Chapter Book, 2022) cites 50 spp. in the Pains karst, only 6 described until the publication of the article, suggesting 44 spp. new undescribed. As the article does not bring a listing by group, here it is not considered to accept these numbers. Simões et al. (Subterranean Biology, 2015) cites 33 troglobic/trogomorphic species in Minas Gerais state, however, without discriminating which are exact troglobics, we decided not to count these taxa. Trevelin et al. (Ecological Indicators, 2019) quotes an incredible 135 possible troglobics in Serra dos Carajás (PA), among Amblypygi (3), Araneae (13), Opiliones (7), Pseudoscorpiona (5), Schizomida (1), Trombidiformes (2), Chilopoda/Scolopendromorpha (1), Diplopoda/Glomedidesmida (3), Diplopoda/Polydesmida (11), Diplopoda/Polyxenida (3), Diplopoda/Spirostreptida (3), Collembola (17), Gastropoda (3), Coleoptera (40), Hemiptera (1), Hymenoptera (1), Isopoda (14), Oligochaeta (3) and Tricladida (2). Considering all the species already described for this region, group by group, in this amount, we would still have a potential minimum of 104 new troglobics, bringing Brazilian diversity to c. 750 spp. However, we do not include this record here due to the lack of a more detailed inventory.
BY GROUP
Largest groups, including non state-endemic species: Isopoda (114), Collembola (88), Diplopoda (66), Araneae (67), Coleoptera (48), Pseudoscorpiona (41), Opiliones (31), Actinopteri (27), Palpigradi (25), Gastropoda (23), Chilopoda (22), Amphipoda (21), Platyhelminthes (15), Hemiptera (12), Amblypygi (11) and Diplura (10).
BY STATE
Only 14 troglobics in Brazil are non states-endemics: one shared with Bolivia; three simultaneously in São Paulo and Paraná; three simultaneously in CE/RN; PA/AP, MG/SP, BA/SE and MS/MT one species each; one occur in MG/SP/PR; one in Minas Gerais, São Paulo and Mato Grosso do Sul; and one in Minas Gerais, São Paulo, Goiás and Paraná - 6 are Pseudoscorpionida.
If you count described and undescribed species state endemics, the numbers are: Bahia (221), Minas Gerais (182), São Paulo (65), Rio Grande do Norte (55), Mato Grosso do Sul (42), Pará (38), Goiás (24), Espírito Santo (11), Tocantins (9), Mato Grosso (4), Paraná (4), Ceará (2), Santa Catarina (2), Rio de Janeiro (2), Piauí (1), Amazonas (1) and Rondonia (1). Sergipe no has endemic troglobics.
If we count all the species (inc. non state endemics), we have, the numbers are Bahia (222), Minas Gerais (186), São Paulo (72), Rio Grande do Norte (58), Mato Grosso do Sul (45), Pará (39), Goiás (25), Espírito Santo (11), Tocantins (9), Paraná (8), Ceará (5), Mato Grosso (5), Santa Catarina (2), Rondonia (2), Rio de Janeiro (2), Amazonas (1), Piauí (1), Amapá (1) and Sergipe (1).
Undescribeds for state: Bahia (120), Minas Gerais (84), Rio Grande do Norte (51, one non-endemic), São Paulo (32), Mato Grosso do Sul (31), Espírito Santo (10), Goiás (10) and Tocantins (6) - 327 spp., all but one (Cirolanidae) state endemics.
BY MUNICIPALITY
Most distributions followed the original descriptions of the species, or local inventories. Another important source consulted was Pires et al. (CBE, 2015) and [1]. 147 municipalities has troglobics in Brazil. The species that was identified in the most municipalities was Trogolaphysa bellinii (Collembola/Paronellidae), having been identified in 10 municipalities in Minas Gerais.
AM ‣ a single species in Presidente Figueredo.
RO ‣ Porto Velho and São Francisco do Guaporé one each.
AP ‣ two municipalities, Amapá and Macapá.
PA ‣ 11 municipalities, Parauapebas (21), Canaã dos Carajás (13), Curionópolis (6), São Felix do Xingu (3), Altamira (2), Medicilândia (3), Ananindeua (1), Belém (1), Benfica (1), Brasil Novo (1), Eldorado dos Carajás (1), Salvaterra (1).
PI ‣ one sp. in Coronel José Dias.
CE ‣ 4 munipalities: Quixerê (2), Santa Quitéria (1), Aiuaba (1), Ubajara (1).
RN ‣ 5 municipalities: Felipe Guerra (41, 38 endemics), Baraúna (9, 8 endemics), Dix-Sept Rosado (9, 7 endemics), Mossoró (2), Apodi (1).
SE ‣ a single species from in Japaratuba and Laranjeiras, also in Bahia.
BA ‣ 28 municialities: Carinhanha (59), Andaraí (38), Iuiu (32), Campo Formoso (27), Malhada (20), Coribe (10), Iraquara (7), Itaetê (6), Santana (6), Serra do Ramalho (6), São Desidério (4), Feira da Mata (5), Morro do Chapéu (3), Palmeiras (3), Paripiranga (3), Varzea Nova (3), Lençóis (2), Santa Maria da Vitória (2), Bom Jesus da Lapa (1), Canálopis (1), Curaçá (1), Ituaçu (1), Maria da Vitória (1), Miranguara (1), Mucugê (1), Nova Redenção (1), Ourolândia (1) and Pau Brasil (1).
TO ‣ 4 municipalities: Dianópolis (6), Aurora de Tocantins (1), Lagoa da Confusão (1) and Natividade (1).
MT ‣ 4 municipalities: Nobres (2), Curvelândia (1), Rosario do Oeste (1) and Diamantino (1).
GO ‣ 5 municipalities: São Domingos (17, 16 exclusive), Posse (3, 2 exclusive), Mambaí (3, 2 exclusive), Guarani de Goiás (1), Nova Roma (1).
MG ‣ 54 municipalities: Cordisburgo (39), Pains (17), Itacarambi (16), Matozinhos (11), Rio Acima (11), Belo Horizonte (10), Itabirito (9), Mariana (8), Arinos (7), Pedro Leopoldo (7), Barão de Cocais (6), Brumadinho (6), Morro do Pilar (6), Nova Lima (6), Sete Lagoas (6), Vazante (6), Conceição do Mato Dentro (5), Lima Duarte (5), Montes Claros (5), Presidente Olegário (5), Caete (4), Luislandia (4), Santa Bárbara (4), Curvelo (3), Lagoa Santa (3), Prudente de Morais (3), Ibiracatu (2), Itambé do Mato Dentro (2), Monjolos (2), Prados (2), São Gonçalo do Rio Abaixo (2), Arcos (1), Capim Branco (1), Carai (1), Catas Altas (1), Dores de Guanhaes (1), Jaiba (1), Jequitaí (1), Juvenilia (1), Moeda (1), Montalvania (1), Ouro Preto (1), Piumhi (1), Presidente Juscelino (1), Rio Pardo de Minas (1), Santa Maria de Itabira (1), São João da Lagoa (1), São João da Lapa (1), São João da Ponte (1), São Roque de Minas (1), São Sebastião do Rio Preto (1), Unaí (1), Vespasiano (1). ES ‣ 5 municipalities: Vargem Alta (5), Ecoporanga (2), Santa Teresa (2), Conceição do Castelo (1), Pedro Calvário (1).
RJ ‣ two municipalities, Cantagalo and Cambuci, one species each.
SP ‣ 11 municipalities, Iporanga (52), Itu (12), Eldorado (3), Itirapina (2), Guapiara (2), Altinopólis (1), Analandia (1), Apiai (1), Ipeuna (1), Ribeirão Grande (1), Ribeira (1).
MS ‣ 5 municipalities: Bonito (24), Bodoquena (23), Porto Murtinho (4), Jardins (5), Corumbá (3).
PR ‣ 5 municipalities: Rio Branco do Sul (3), Adrianópolis (2), Almirante Tamandaré (1), Dr. Ulysses (1), Ponta Grossa (1).
SC ‣ a single municipality Botuverá (2).
Largest diversities by municipalities (all species): Carinhanha (59), Iporanga (52), Felipe Guerra (41), Cordisburgo (39), Andaraí (38), Iuiu (32), Campo Formoso (27), Bonito (24), Bodoquena (23), Parauapebas (21), Malhada (20), Pains (17), São Domingos (17), Itacarambi (16), Canaã dos Carajás (13), Itu (12), Matozinhos (11), Rio Acima (11), Belo Horizonte (10), Coribe (10), Baraúna (9), Governador Dix-Sept Rosado (9), Itabirito (9), Mariana (8), Arinos (7), Iraquara (7), Pedro Leopoldo (7), Barão de Cocais (6), Brumadinho (6), Curionópolis (6), Dianópolis (6), Itaetê (6), Morro do Pilar (6), Nova Lima (6), Santana (6), Serra do Ramalho (6), Sete Lagoas (6), Vazante (6), Conceição do Mato Dentro (5), Feira da Mata (5), Jardins (5), Lima Duarte (5), Montes Claros (5), Presidente Olegário (5), Vargem Alta (5), Caete (4), Luislandia (4), Porto Murtinho (4), Santa Bárbara (4), São Desidério (4), Caete (3), Corumbá (3), Curvelo (3), Eldorado (3), Lagoa Santa (3), Mambaí (3), Morro do Chapéu (3), Palmeiras (3), Paripiranga (3), Posse (3), Prudente de Morais (3), Rio Branco do Sul (3), São Felix do Xingu (3), Varzea Nova (3), Adrianópolis (2), Altamira (2), Botuvera (2), Ecoporanga (2), Guapiara (2), Ibiracatu (2), Itambé do Mato Dentro (2), Itirapina (2), Lençóis (2), Medicilândia (2), Monjolos (2), Mossoró (2), Nobres (2), Prados (2), Quixerê (2), Santa Maria da Vitória (2), Santa Teresa (2), São Gonçalo do Rio Abaixo (2), Aiuaba (1), Almirante Tamandaré (1), Altinopólis (1), Amapá (1), Analandia (1), Ananindeua (1), Apiai (1), Apodi (1), Arcos (1), Aurora de Tocantins (1), Belém (1), Benfica (1), Bom Jesus da Lapa (1), Brasil Novo (1), Canálopis (1), Cantagalo (1), Cambuci (1), Conceição do Castelo (1), Capim Branco (1), Caraí (1), Catas Altas (1), Coronel José Dias (1), Curaçá (1), Curvelândia (1), Diamantino (1), Dr. Ulysses (1), Dores de Guanhaes (1), Eldorado dos Carajás (1), Guarani de Goiás (1), Ipeuna (1), Ituaçu (1), Jaiba (1), Japaratuba (1), Jequitaí (1), Juvenilia (1), Lagoa da Confusão (1), Laranjeiras (1), Macapá (1), Maria da Vitória (1), Miranguara (1), Moeda (1), Montalvania (1), Mucugê (1), Natividade (1), Nova Redenção (1), Nova Roma (1), Ouro Preto (1), Ourolândia (1), Pau Brasil (1), Pedro Canário (1), Piumhi (1), Ponta Grossa (1), Porto Velho (1), Presidente Figueredo (1), Presidente Juscelino (1), Ribeirão Grande (1), Ribeira (1), Rio Pardo de Minas (1), Rosario do Oeste (1), Salvaterra (1), Santa Quitéria (1), Santa Maria de Itabira (1), São Francsco do Guaporé (1), São João da Lagoa (1), São João da Lapa (1), São João da Ponte (1), São Roque de Minas (1), Ubajara (1), Unaí (1) and Vespasiano (1).
BRAZILIAN TROGLOBIC CAVE (BLACK DOTS) AND PHREATOBIC (RED POINTS, INCLUDING BOLIVIAN RECORD OF PHREATOBIUS SANGUIJUELA) FOR BRAZIL. EACH POINT MAYBE SEVERAL CAVES/MUNICIPALITIES.
OTHER NOTES
Many links of Brazilian cave fauna was found in Laboratório de Estudos Subterrâneos/UFSCar - São Carlos. An important socio-environmental description of a Brazilian cave environment refers to the Pains karst, in Minas Gerais (Ferreira, Zampulo & Souza-Silva, CHAPTER BOOK, 2020).
Important note: no photos used on this page of troglobics species belong to the author of this site. All were taken from the internet. If any author wants it removed, we will do so as soon as possible.
Of all the phyla that evolved troglobic species, Brazil has representatives of all except Cnidaria and Nematoda. The vast majority of infra-phylum lineages are found in the country, with the exception of Themnocephalida (S Europe and Papua New Guinea), Prorhyhchidae (South Africa), Sabellida (NE Italy, Slovenia, Croatia, Bosnia & Hercegovina), Phyllodocida (SE Mexico and Caribbean, Canary Is., Oman, Socotra, W. Australia, Papua New Guinea, New Zealand and Fiji), Branchiopoda (Spain, Romania, France, Slovenia, Bosnia and Herzegovina), Remipedia (Mexico, Caribbean, Canary Is. and NW Australia), Leptostraca (Turks y Caicos), Anaspidaceae (Australia and Tasmania), Thermosbaenacea (widely worldwide), Mysida (several places worldwide), Mictacea (Bermudas), Bochusacea (Bahamas), Tanaidacea (Bermuda and several islands of the South Pacific), Cumacea (Bermudas, Bahamas and Jamaica), Decapoda/Stenopodidae (Bahamas), Decapoda/Gebiidea (Bahamas), Decapoda/Caridea (widely worldwide), Decapoda/Astacida (Papua New Guinea, Cuba, Mexico, North America), Decapoda/Brachyura (widely worldwide), Collembola/Neelipleona (Europe and Asia), Insecta/Grylloblattoidea (unvailable range), Insecta/Psocoptera (Venezuela and Old World), Insecta/Lepidoptera (Philippines and Hawaii), Arachnida/Opilioacarida (Cuba, Thailand and Vietnam), Arachnida/Schizomida (Cuba, Jamaica, Belize, Mexico, U.S.A., Ecuador, and Australia), Arachnida/Thelyphonida (possibly in Laos), Arachnida/Ricinulei (Venezuela, Belize, Cuba and Mexico), and Amphibia (U.S.A. and Balkans in Europe).
1 PHYLUM PORIFERA ‣ [2] lists only one cave sponge in World; however, there are three described species, all in Spongillidae, obligatory subterranean: Eunapius subterraneus Sket & Velikonja in Ogulin caves, Croatia, and two in Brazil: Arinosaster patriciae Volkmer-Ribeiro, Tavares-Frigo, Ribeiro & Bichuette, collectd in Parecis System, Mato Grosso state (Neotropical Biology and Conservation, 2021); and Racekiela cavernicola Volkmer-Ribeiro, Bichuette & Machado, collected in Lapa dos Brejões, northern Bahia state (Volker et al., Neotropical Biology and Conservation, 2010). Both Brazilian troglobic species cited here are corroborated by [6]. A fourth species, undescribed, was discovery in center Mexico (Legendre et al., Subterranean Biology, 2023).
2 PHYLUM CNIDARIA ‣ many records of Cnidaria in freshwater caves can be found in Zagmaster et al. (Speleobiology, 2011), Mexico, U.S.A., Australia and several center European countries. Velkovrhia enigmatica Matjasic & Sket, 1971 (Bougainvilliidae, Hydrozoa, PHOTO), from Dinarides in the Balkan Peninsula, is the only troglobic freshwater Cnidarian. It has been known from five caves in Slovenia (3), Croatia (1) and Bosnia (1), by Magmajster (Natura Sloveniae, 2003). Infortunely, it is not listed in [3] among freshwater Cnidaria.
3 PHYLUM PLATYHELMINTHES ‣ by our survey, 4 groups of flatworms have troglobites: Alloeocoela (1, Prorhynchidae, Geocentrophora in U.S.A.) in [2], Tricladida, Proseriata (1, in South Africa, SEE) and Temnocephalida.
In Tricladida and Temnocephalida, for Western Europe, there is consolidated data only for the Balkans [29], and a generic mention [21], which consolidates 111 troglobites for the region in this group (99 Tricladida and 12 Temnocephalida). Tricladida also are reported elsewhrere: 3 in Africa [7], 4 in former USSR except Baltic countries [21], 32 in U.S.A./Canada [2], 1 in Jamaica [24], 6 in Mexico and 15 in Brazil. In these numbers, our survey will reliably point to at least 173 known species of troglobitic Platyhelminthes. Record of a Temnocephalida for a crab in Papua New Guinea as troglobic is here rejected [22]. Troglobics absents in China [2].
All Brazilian troglobic Platyhelminthes belongs order Tricladida [1] in all three clades: Continenticola (12 in Girardia, Dugesiidae), Maricola (1) and Cavernicola (2). Four undescribed spp. was cited from Mato Grosso do Sul [MS/1], São Paulo [SP/2], these Continenticola, and one Cavernicola for Rio Grande do Norte [RN/1], and one of undetermined order for Minas Gerais [MG/7]. Brazil is the only country in the world with troglobic representatives in the three groups of Tricladida. Sluysia triapertura Leal-Zanchet & Souza, 2018 represents the first Maricola triclad living in freshwater within a cave (Souza et al., Invertebrates Systematics, 2018).
UNDETERMINED TURBELLARIA
CAVERNICOLA
CONTINENTICOLA
MARICOLA
4 PHYLUM NEMATODA ‣ despite the extreme diversity of habitats they can occupy in contrast with the extreme morphological homogeneity, there is no systematic culture about cave nematodes and even criteria to define whether a species is troglobotic do not exist, with a large amount of collections but little precision of what is in fact a Nematoda troglobic. Carefully consulting the Du Preez (Nematology, 2017), only seven spp. possibly are true troglobics: Desmoscolex aquaedulcis Stammer, 1935 (Slovenia), Thalassoalaimus aquaedulcis Schneider, 1940 (Slovenia), Halalaimus stammeri Schneider, 1940 (Slovenia), Hemicycliophora aquatica Loos, 1948 (Belgium), Stenonchulus troglodytes Schneider, 1940 (Slovenia, Austria), Mylonchulus cavensis Schneider, 1940 (Hungary), and Chronogaster troglodytes Poinar & Sarbu, 1994 (Romania). Axonchium sbordonii Zullini, 1973, from Sima del Ojito, Chiapas, Mexico, is rejected in this work (Mexican Cave Fauna, 2022). In adition, one possibly troglobic species, in Rhabditida, was collected in Colombia [19 + notes].
5 PHYLUM NEMERTEA ‣ by [2], 3 of all 20 freshwater Nemertea are possibly troglobics (stygobitics): Prostoma puteale Beauchamp, 1932 in France, Switzerland, and Germany, and Prostoma hercegovinense Tarman, 1961 in Bosnia and Herzegovina, but one species has been described from New Zealand also. In addition, Brazil has two possibly troglobic Nemertea, one in Alto do Ribeira region in São Paulo state [SP/2], another from C Minas Gerais state [MG/1]. If confirmed, they will be the first troglobic Nemertea described for the New World.
6 PHYLUM ANELLIDA ‣ based on our surveys, three clades within Annelida has troglobics: Sabellida, Phyllodocida and Clitellata. Some families have adapted to living in the water column of marine/anchialine caves, including the normally benthic Protodrilidae, Nerillidae, and Scalibregmatidae. The aptly named genus Troglochaetus (Nerillidae) is also found in the hyporheic zone of caves, wells and springs, with T. beranecki Delachaux, 1921 currently thought to occur in both Europe and North America. Troglochaetus has rudimentary parapodial cirri and, like all nerillids, development is direct, either via anexternal brood or in cocoons (Galsby et al., Diversity, 2021).
Numerous marine species live in conditions similar to cave conditions: inside the bottom material, under stones and in the rock fissures. In submarine (anchihialine) caves, stygobiotic forms very tolerant to changes of environmental conditions were found. In Central and South America, rich polychaete fauna was found in cenotes having the connection with the ocean. Morphological adaptations to cave environment such as the absence of body pigmentation and lack of eyes are rarely observed in polychaetes; principally therefore they can be classified as stygobionts only on the basis of their habitat [2].
Our survey point to 163 known troglobic Annelida, 148 in Clitellata, one in Sabellida, and 14 in Nemanereis.
SABELLIDA
Sabellida has one full troglobic, Marifugia cavatica Absolon & Hrabe, 1930 (Serpulidae), a eyeless species known from caves situated in Dinaric Karst (northeastern Italy, Slovenia, Croatia, and Bosnia & Hercegovina), but occurs in small, isolated areas, considered to be a Tertiary relict of marine origin. Even though some localities are situated few kilometers from the seashore this species has never been found in brackish or salt water. Individuals live in calcareous tubes which are attached to the walls, roofs, and bottoms of water-filled corridors forming colonies of various dimensions. In the last years, live animals were collected in a few caves only, whereas aggregations of their broken tubes were found in many localities [2].
PHYLLODOCIDA
Phyllodocida has at least 14 cavernicolous (however, all inconcluse be troglobics) species representing genus Namanereis (Nereididae). C. J. Glasby et al. (ZJLS, 2014) lists all stygobitics Namanereis: N. cavernicola Solís-Weiss & Espinasa, 1991 in SE Mexico and Caribbean (some populations extreme continental, found in cave pools above 1600 m asl and over 170 km from the coastline), two undescribeds from Canary Is., 5 endemics to Caribbean, N. araps Glasby, 1997, in Oman, two in Socotra, N. pilbarensis Glasby, Fiege & Van Damme, 2014 in W. Australia, N. beroni Hartmann-Schröder & Marinov, 1977 in Papua New Guinea and N. tiriteae Winterbourn, 1969 in New Zealand and Fiji.
CLITELLATA
Small troglobic Clitellata live among the grains of sand and gravel saturated with water (interstitial waters) along streams and in cave stagnant water bodies. Bigger ones 'are motile on on the bottom surface or swim using setae as locomotory organs. Naidids and lumbriculids generally are anchored inside the sediments with the anterior part of the body, waving the posterior end for better oxygenation of the surrounding water. They move slowly when the food is used up or environmental conditions deteriorate, that is when oxygenation decreases or products of metabolism accumulate. Like nematodes, some stygobiotic Clitellata species can form cysts, which help them to survive in small, periodically desiccated cave pools. The majority of oligochaetes feed on organic matter from sediments or soil. Only a few stygobiotic species are predators. [2]. Aquatic members have a habitus that makes them pre-adapted to live in the subterranean environment. In addition, they do not exhibit any peculiar morphological adaptations to subterranean life that can be seen in other subterranean organisms (loss ofeyes, elongation of appendages and body, loss of pigmentation, increase in sensory struc-ture). Therefore, stygobitic nature of the species can only be inferred from their exclusive presence in the subterranean environment. Despite their morphological pre-adaptation, few aquatic Clitellata are exclusively present in groundwater.
Our survey listed 148 Clitellata, with 112 mentioned in [20], 9 listed in [21], 4 news for U.S.A. after [25], 6 adds for Brazil (all, free and parasitics), two free-Clitellata adds for Mexico, 8 Branchiobdellida for U.S.A. and Mexico, and 7 Hirudina except Brazilian and Georgian news. By countries, are (5:)89 in Europe [20], 18 in U.S.A./Canada (8 in [20], 4 news free-Clitella, six Cambarincola), 10 in former USSR (all by [21], one leeche shered with Europe), 4 in Mexico (2 free-life, two parasitics), six in Brazil, 3 in Africa [20], one in Iran (Hirudina), one in China (Hirudina), 2 in Ecuador [20], two in Nippon [20], one in New Zealand [20], two in Oman [20], one in Australia [20], 5 in Cuba [20], three in Caribbean except Cuba [20] and one in Venezuela [20].
Enchytraeidae: (6/)12 troglobic spp. in caves, 10 in Europe (Poland, Hungary, Romania, Greece) and two in Ecuador: Marionina ecuadoriensis Righi and Hemienchytraeus mauriliae Righi.
Haplotaxidae: (3/)9 troglobic spp., 8 cited in [20] in Europe (5, Spain, France and Bulgary), Nippon (1) and Guinea (2), and one added for U.S.A. in [25].
Lumbricidae: (2/)2 spp. (Eophila hypogea Malevics, 1947 and Archaeodrilus cavaticus Czerniavsky, 1880), cited for Georgia and Abkhazia [21].
Lumbriculidae (includes Dorydrillidae): by [20], (9/)37 troglobic spp., mainly in Europe (UK, Spain, France, Belgium, Germany, Switzerland, Italy, Estonia, Slovenia, Czech Republic, Croatia, Serbia, Bosnia-Herzegovina, Macedonia, Romania), one in Japan and 7 in U.S.A., in Virginia, Alabama, Tennessee in E, Oregon and California in W: Spelaedrilus multiporus Cook, Stylodrilus beattiei Cook, S. californianus Rodriguez, S. wahkeenensis Rodriguez & Coates, Tenagodrilus musculus Eckroth & Brinkhurst, Trichodrilus allegheniensis Cook, and T. culveri Cook. Additionally, undescribed species, counted here as only one, are known from Abkhazia.
Parvidrilidae: two troglobic spp. in Parvidrilus, one in Italy and Slovenia, another in Alabama, U.S.A. (SEE).
Phreodrilidae: (3/)5 troglobic spp., in New Zealand (1), Oman (2), Morocco (1) and Australia (1).
Tubificidae: (19/)48 troglobics spp., nine spp. in New World: Phallodriloides macmasterae Erséus, 1986 in Bermuda, four Pristina and Clitellio cavernicolus Botea, 1983 in Cuba, Dero haitiensis Dumnicka, 1986 and Spirospermoides stocki Dumnicka 1983 in Haiti and adjacent islands, and Krenedrilus papillatus Dumnicka, 1983 in Venezuela, 39 in Europe (Portugal, Spain, France, Germany, Switzerland, Italy, Poland, Czech Republic, Slovenia, Macedonia, Greece and Turkyie; Gianius aquaedulcis Hrabě, 1960 rejected in U.S.A./New York, SEE). Additionally, [21] includes 5 more species, from Russia, Uzbekisthan, Abkhazia and Georgia.
Hirudina/Hirundinina: at least 10 troglobics species in three families (6 in Europe and former USSR, 1 in Iran, 1 in China, 2 in Brazil), Haemopidae, Haemadipsidae and Erpobdellidae, usually exhibiting two specific morphological features, namely milky-white coloration and lack of eyes or eye pigment (Cichocka et al., Zootaxa, 2015). Troglobics leeches cited in literature fall in this description: Haemopis caeca Klemm & Sarbu, 1998 from Dobrogea in Romania (Haemopidae), Erpobdella absoloni Johansson, 1913 (sometimes as Dina absoloni) in N Italy–Balkans–Turkey–Georgia [21], Erpobdella underminated spp. from Abkhazia [21], E. borisi Cichocka & Bielecki from NW Iran, E. krasensis Sket, 1968, and E. mestrovi Kerovec, Kučinić & Jalžić, 1999 from Croatia (Cichoka, 2015), with possibly undescribed species, probably Erpobdellidae, in caves of China and the U.S.A. (Sket & Trontelj, Hydrobiologia, 2006), non counted here. Two known species of Haemadipsidae, Leiobdella jawarerensis Richardson, 1974 and Haemadipsa cavatuses Yan get al., 2009 possess five pairs of eyes, which is the only pigmented part on its body, inhabiting the aphotic zone of a caves in Papua New Guinea and China, respectively (Cichoka, 2015). However, by [22], papuan species is no accepted here as troglobic. Two species of two diferent families are troglobics in China. Two possibly undescribed troglobics species was collected in C Minas Gerais state [MG/1] and [MG/7], Brazil. If confirmed, they will be the firsts troglobics Hirudinea described for the New World.
Hirudina/Branchiobdellida (crayfishworms) are also known from caves as stygobitic; they are usually ectoparasites of crustaceans, some of them (similarly to flatworms Themnocephalida) were found on one to three host species only [2]. At least 8 troglobics, two in Mexico (Mexican Cave Fauna, 2022) and six in U.S.A. [2].
7.1 MOLLUSCA/BIVALVIA ‣ typical troglobic mollusks usually exhibits the following characteristics: (1) absent or reduced pigmentation, (2) absent or reduced eyes, (3) some reduction in shell size (reduction in size compared to their epigean relatives). The two classes with non-marine species have troglobic species. Bivalvia includes six troglobics described worldwide: 4 Euglessa (Euglessidae) from East Europe [21], in Russia (1), Abkhazia (2) and Georgia (1); Congeria kusceri Bole, 1962 (Dreissenidae, Total Croatia News), known only from Slovenia to SW Bosnia and S Dalmatia; and Eupera troglobia Simone & R. L. Ferreira, 2022 (Sphaeridae), known only W Tocantins state, in center Brazil, unique troglobic bivalvia in Western Hemisphere.
7.2 MOLLUSCA/GASTROPODA ‣ troglobitic (terrestrial) snails are all pulmonates, except for the prosobranch Pholeoteras euthrix Sturany, 1904, known from caves in southern Croatia, southern Herzegovina and Greece (ANIMALBASE). All stygobitic Gastropoda are minute prosobranchs with shells smaller than 2mm except Acroloxus tetensi Kušcer, 1832 from the Dinaric Mountains in Slovenia, and Hydrophrea from New Zealand, which are pulmonates.
There is no exact data on the diversity of troglobic Gastropoda in Western Europe. Here, we will adopt artifically the number of 368, which is the sum of 143 stygobites from France [2], 169 stygobites from the Balkans (Falniowsk, A. et al., Hydrobiologia, 2021), and 56 troglobites mentioned in [2]. Europe is home to 11 genera of strictly troglobitic snails: Troglaegopis, Sciocochlea, Speleodentorcula, and Spelaeoconcha (all monotypics), Paraegopis (5, 1 TB), Argna (6, 1 TB), Klemmia (2, 1 TB), Cryptazeca (4, 2 TB), Virpazaria (7, 3 TB), Spelaeodiscus (10, 9 TB) and Zospeum (35 TB), mainly in the Dinaric karst.
Troglobic (both stygobitic and troglibiont) Gastropod are found elsewhere, 16 in Africa (15 Litthorimorpha, 1 Stylomattophora), 22 in former USSR except Baltic countries [21], 1 in Iran [8/9/10], 2 in China [2, both Pulmonata, in two families], in New Guinea [22], 4 in Australia [12], 1 in New Zealand [2, Hydrophrea], 32 in U.S.A./Canada [2], 9 in Mexico [18, all stygobitics] and 23 in Brazil. In these numbers, our survey will reliably point to at least 479 known species of troglobitic Gastropoda. Recent data point to (5:20/)39 spp. for aquatic members in USA/Mexico, by Gladstone et al. (Conservation Biology, 2022).
Cavallaria et al. (Zoologia, 2022) cites 19 described troglobic gastropoda from Brazil, however, troglobic status for many of them are rejected by several autors. Here we accept 23 names as troglobics in Brazil, 6 already described and mentioned expressly in R.L.Ferreira et al. (Zoologia, 2023), two in Salvador & Bichuette (Zoosystematics and Evolution, 2024), 14 not yet described, six Potamolithus in São Paulo [SP/1], two in Mato Grosso do Sul [MS/1] and six in Bahia [BA/2][BA/3][BA/4], and three described latter. Overall, Brazil has at least 9 genera and 9 families.
UNDETERMINED FAMILIES
CAENOGASTROPODA
DIPLOMMATINIDAE
EUPULMONATA
CHAROPIDAE
COCHLIOPIDAE
GASTROCOPTIDAE
LITHOGLYPHIDAE
POMATIOPSIDAE
SCOLODONTIDAE
SYSTROPHIIDAE
TOMICHIIDAE
8 PHYLUM ONYCHOPHORA ‣ by [4], officially only two trogobitic Onychophora are known: Peripatopsis alba Lawrence, 1931 (Peripatopsidae) of South Africa and Speleoperipatus spelaeus Peck, 1975 (Peripatidae) from Jamaica [24], with probable species also in the Galapagos. Unofficially, there are one troglobic species also in Brazil, not yet formally described, in caves of the Bodoquena System, in Mato Grosso do Sul state, and may even be a new genus [MS/1].
PERIPATIDAE
Arthropoda includes 17 lineages, 14 has troglobics: Aracnida, Chilopoda, Pauropoda, Symphyla, Diplopoda, Oligostraca (former Ostracoda), Thecostraca, Copepoda, Branchiopoda, Remipedia, Malacostraca, Collembola, Diplura and Insecta. Exceptions: Pycnogonyda (marine), Cephalocarida (marine) and Protura.
Former Crusteca is divided in eight class-like levels nowdays. Nominally, 26 lineages includes troglobics, in all three types: terrestrial, freshwaters and marine.
SINGLE HABITAT
All troglobics in Branchipoda/Diplostraca (Europe), Anaspidacea (Australia, New Zealand, Argentina and Chile) and Astacidea (40, PNG, U.S.A., Mexico, Cuba) are fully freshwaters and unknown in Brazil.
All troglobics in Spelaeogriphacea and Bathynellacea are fully freshwaters, both with troglobic members in Brazil.
All troglobics in Stygiomysida, Thecostraca/Tantulocarida (1, Canary Is.), Cumacea, Tanaidacea, Remipedia (25, Mexico, Belize, Caribbean, Canary Is., Australia), Stenopodida (1, Bahamas), Gebiida (1, Bahamas), Mictacea (1, Bermuda), Bochusacea (1, Bahamas) and Leptostraca (1, Turks y Caicos) are fully anchialine or marine halotroglobic, unknown in Brazil.
MIXED HABITAT
Anomura includes troglobics in saltwaters of Canary Is., and freshwaters in SE Brazil.
Marine troglobics in Amphipoda, Isopoda, Copepoda, Ostracoda and Anomura are unknown in Brazil, besides troglobic members in freshwater caves inland.
Neither freshwater or marine troglobics in Mysida, Thermosbaena, Caridea or Brachyura are known in Brazil.
9.1 ARTHROPODA/ARACHNIDA ‣ all orders of Archnida has troglobics except Holothrida and Solifugae. 337 troglobic Arachnida occur in North America (in Acari, Pseudoscorpiones, Araneae, Opiliones, Schizomida, Scorpionida), 144 in China [2] and 184 in Mexico [5], and 178 in Brazil.
Among Acari, two superorders and four orders contain troglobic species: Parasitiformes with the orders Opilioacarida and Gamasida, and Acariformes with the orders Sarcoptiformes and Trombidiformes. U.S.A. has (10:17/)26 spp. troglobic Acari, 13 in Rhagidiidae [2]. Despite the extreme diversity of mites in caves in Brazil, with 126 families collected in caves in the country [6], only one spp. is formally described, and another 7 are identified as troglobic without description, in Bahia [BA/1][BA/3], Minas Gerais [MG/1][MG/4] and Mato Grosso do Sul [MS/1].
A. OPILIOACARIDA ‣ none Brazilian species is true troglobics, despites several cave collections (Berbardi & Borges-Filho, Subterranean Biology, 2018). Troglobics occur only in Cuba, Thailand and Vietnam [4].
B. IXODIDA ‣ by Galán & Herrera (BSVE, 2006), Antricola silvai Cerny, 1967 from Venezuela maybe a troglobic. If confirmed, it will be the only record of an Ixodida troglobic worldwide. Huge populations occur in caves in Brazil, associated with bats, all troglophiles.
C. MESOSTIGMATA ‣ few informations. In this order, a large number of cave species are permanent or temporary ectoparasites of bats, but most are free living. Three troglobic undescribed species in Brazil, collected in Minas Gerais [MG/1] and Bahia [BA/3] states.
UNPLACED FAMILY
DITHINOZERCONIDAE
PACHYLAEPIDAE
D. TROMBIDIFORMES ‣ Prostigmata (more than 20,000 spp.), includes a few poorly known cave species in the families Leeuwenhoekiidae, Trombiculidae, and Trombidiidae, frequent on or close to guano accumulations intropical caves, and bat or invertebrate parasites at larval stage. Family Proterorhagidiidae has a single relictual troglobic, Proterorhagia oztotloica Lindquist & Palacios-Vargas, 1991 from Mexico. Troglobic Rhagidiidae are often linked to cold caves in the Holarctic [4]. One troglobic Rhagidiidae also occur in Mexico. In Brazil occur a single described species and two undescribeds in Bahia [BA/1] and Minas Gerais [MG/4]. Venezuela has eight troglobic Hydracnidia in three families (Galán & Herrera, BSVE, 2006).
ERYTHRAEIDAE
RHAGIDIIDAE
