UPDATED IN 25∙04∙2024
1 INTRODUCTION
For the purposes of this text, based on Trajano & Bichuette (BOOK, 2006), we will use the following terminology: (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). The entire text of this article refers only to troglobionts, including aquatic and terrestrial forms under the same term.
Troglobites 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.
2 REFERENCES
[1]: Gallão & Bichuette (ZooKeys, 2018) - brazilian primarily troglobite list; [2]: White & Culver (Enciclopedia of Caves, 2019, 3th edition) - diversity overall in caves; [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).
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single publication; however, due to the fact that it is a book written with several contributions, many data are still out of date and others are conflicting with others from the same work.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.
3. WORLD DIVERSITY AND NUMBERS
10 phylla has true troglobitic/stygobitic known for science. Although there is no consolidated data, our work believes that c. 9,650 obligatory subterranean species occur in the world: approximately 5,000 troglobites from Europe [2, pg. 136], 1,200 had been described from Asia [2, pg. 136], 500 from Africa [2, pg. 136], c. 1,850 from North America ([2, pg. 136], Palacios y Vargas (UMAE, 2013)), c. 700 for South America (this source, inc. undescribeds from Brazil, c. 500) and c. 400 in Australia (M. T. Guzik et al., Invertebrate Systematics, 2010). If we estimate minimally realistically the estimative of M. T. Guzik et al. (2010), with c. 4,140 troglobitic/stygobitic spp. from Australia, would be, worldwide, an estimate of 13,390 spp., only 4% in Brazil.
Of all the phyla that evolved troglobiont 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 one at a crab in Papua New Guinea), Prorhyhchidae (South Africa) of Platyhelminthes; Sedentaria/Sabellida (NE Italy, Slovenia, Croatia, Bosnia, and Hercegovina) and Errantia/Phyllodocida (SE Mexico and Caribbean, Canary Is., Oman, Socotra, W. Australia, Papua New Guinea, New Zealand and Fiji) in Annelida; Branchiopoda (Spain, Romania, France, Slovenia, Bosnia and Herzegovina), Remipedia (Mexico, Caribbean, Canary Is., and in 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/Astacidae (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), in Arthropoda; and Amphibia (U.S.A. and Balkans in Europe).
3 NOTES BY COUNTRIES
For publications about on extremely diverse caves around the world, see Diversity: Special Issue 'Hotspots of Subterranean Biodiversity'.
U.S.A./CANADA
By [2], the current number of described troglobits from the U.S.A. and Canada, which includes Alaska and Hawaii, is 1,353 spp., representing 5 phyla, 15 classes, 51 orders, 124 families, and 288 genera (1/2 are spiders, diplopods, pseudoscorpiones or beetles, and 1/5 amphipodes or isopods). U.S.A. and Canada, joined, has obligatory subterranean species in several groups without formal description in Brazil: Cestoda (1), Clitellata (17), Maxillopoda (17), Ostracoda (16), Bathynellacea (19), Thermosbaenacea (1), and Schizomida (2). However, considering also undescribed species, the list is shorter: Cestoda (1), Thermosbaenacea (1) and Schizomida (2).
Excepting Amphipoda, Coleoptera, Diplopoda and Pseudoscorpiona, the North American advantage over Brazil drops 585, to just 221 spp.
CUBA
Cuba has 112 troglobitic species, all Arthropoda, 52 Crustaceae, 33 Aracnida, 2 Myriapoda and 21 insects, especifically: Remipedia (1), Ostracoda (5), Copepoda (18), Mysidacea (5), Isopoda (7), Decapoda (13), Amphipoda (3), Scorpiones (1), Pseudoscorpiones (3), Amblypygi (1), Schizomida (5), Ricinulei (4), Opiliones (4), Araneae (7), Acari (12), Zygentoma (3), Collembola (7), Orthoptera (4), Blattoidea (1), Coleoptera (6), by Díaz et al. (Biodiversidad de Cuba, cap. 15, 2015).
CHINA
China has 742 spp. of obligate subterranean species, more than a half are fishes, beetles and spiders [2].
PERU
Only 3 spp. in Peru the following can be considered troglobites: the collembolan Acheroxenylla lipsae Palacios-Vargas, 2020 (Collembola, Poduromorpha, Hypogastruridae), the catfish Astroblepus riberae (Cardona and Guerao, 1994) (Actinopterygii, Siluriformes, Astroblepidae) and Caecopactes chullachaqui Campos-Filho, Sfenthourakis and Bichuette (Isopoda, Scleropactidae) are known (Campos-Filho et al., Nauplius, 2023).
COLOMBIA
Angarita-Sierra (CCE/2018) cites 56 troglobitics in Colombia: two Annelida (Haplotaxide, Enchytraeidae one each), Tityus grottoedensis (Scorpionida), seven Opiliona (in seven families), two Amblypygy (Phrynidae), 16 spiders (in 11 families, Barychelidae, Araneidae, Anyphaenidae, Lycosidae, Tetragnathidae, Pisauridae without Brazilian troglobitics), two Chilopoda (undescribed Scolocryptops, Psilliodes gaildingii), one Isopoda (Armadillidium sp.), 5 Coleoptera (two in Staphylinidae; Carabidae, Nitidulidae, Passalidae one each), two Ephemeroptera (if confirmed, will be the firsts troglobitcs in this order woridwide), one Blattoidea (Blaberidae), one Dermaptera, two Diptera (Drosophilidae and Mycetoohilidae one each), three Hemiptera (Cydnidae, Cavernicola pilosa in Reduviidae, Strudivelia cinctipes in Veliidae), one Lepidoptera, two Trichoptera (in Hydrobiosidae and Hydropsychidae), 7 pisces (all Trichomycterus) and Steatornis caripenis. Apart this work, Jorottui (Amblypygy) was recently described for the country. In addition to these 57 spp., Angarita-Sierra et al (Chapter Book, 2019) cites as troglobitics in Colombia a Rhabditida (Nematoda), one more Pselliodes (Chilopoda) and more three Diplopoda.
Steatornis caripenis assignated in work a troglobitic is a obviously a huge error. Troglobitic 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), 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). With these exceptions, Colombia has, with some consistency, 37 spp. of troglobites. However, troglobitic status of Nematoda, both Ephemeroptera, Dermaptera, both Diptera, Lepidoptera and both Trichoptera need strong validation to be fully credited.
VENEZUELA
The most recent listing for Venezuelan troglofauna dates back to 1 and a half decades: Galán & Herrera (Boletín de la Sociedad Venezolan, Espeleologia-BSVE, 2006). There are 46 troglobites listed for the country: nine Acari (4 families, none with troglobitics in Brazil), four Opiliona (all in Trinella, Agoristenidae), two Amblypygy (both Charinus), one Scorpionida (Taurepania trezii, 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), seven fishes (in 4 families). Cyathura univam (Isopoda) and Metaniphargus venezuelanus (Amphipoda) are anchialines.
SOUTH AMERICA EXCEPT BRAZIL, VENEZUELA, COLOMBIA AND PERU
See Almanaque Z post.
MEXICO
Mexico has 528 spp. of true troglobitics/stygobitics in the following groups: Platyhelminthes (six spp. in two genera, 4 Continenticola, 2 Cavernicola), Annelida (5 spp. in 3 genera, 1 Errantia, 2 Oligochaeta, 2 Branchibdellida), Gastropoda (9 spp. in 5 genera of 3 families), Remipedia (2, both in Xibalbanus), Copepoda (12 spp. in 8 genera at 5 families), Ostracoda (4, three genera of three families in two orders), Amphipoda (22 spp., two families, Bogidiellidae and Hadziidae), Thermosbaenacea (a single species, Tulumella unidens from Quintana Roo), Mysida (7 spp. in three genera at 3 families), Decapoda (27 spp.: 17 shrimps in six families, 5 crayfishes at Cambaridae, and 5 crabs in two families), Isopoda (52 spp. in 11 families), Collembola (42 spp. in nine families), Diplura (8, two Litocampa, two Juxtlacampa, one Oncinocampa and three Tachycampa), Zygentoma (13 spp. in three genera), Orthoptera (six spp., all in Phalangospsidae), Coleoptera (44 spp., Carabidae/30, Dytiscidae/1, Histeridae/4, Leiodidae/8, and Ptinidae/1), Hemiptera (3, all Fulguromorpha), Diplopoda (73, in 15 families at six orders), Chilopoda (5, two Lithobiidae, 3 in Newportia), Trombidiformes (2), Pseudoscorpiona (40 spp. in 5 families), Araneae (59 spp. in 11 families), Opiliona (16 spp. in 4 families), Amblypigy (11, all in Phrynidae), Schizomida (23, 14 in Hubbardiidae, 9 in Protoschizomidae), Ricinulei (11, all in Pseudocellus), Scorpionida (13 spp. in 8 genera at 4 families), and Actinopterigii (12 spp. in six genera and 5 families at 5 orders), by Almanaque Z (SEE).
Recently, a non described troglobotic sponge was discovery in Mexico (Legendre et al., Subterranean Biology, 2023).
Mexico no has described troglobitic in Nematoda, Nemertea, Onychophora, Mollusca Bivalvia, Bathynellacea, Spelaeogriphacea, Blattodea, Dermaptera, Diptera, Hymenoptera, Pauropoda, Symphyla, Palpigradi, and Amphibia. Brazil has more species than Mexico in all groups except in Remipedia, Copepoda, Ostracoda, Amphipoda, Thermosbaena, Mysida, Decapoda, Zygentoma, Coleoptera, Diplopoda, Pseudoscorpionida, Araneae, Amblypigy, Schizomida, Ricinulei, and Scorpionida, all in Arthropoda.
TROGLOBITIC DIVERSIEST WORLD CAVE SYSTEMS
4 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.
A TABLE OF ALL NUMBERS USED IN THIS POST
For species tagged from Brazil, this article lists in cards 650 spp. (all endemics except Phreatobius sanguijuela), 142 valid species described in [1] plus:
:: [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); and Ochyrocera ibitipoca (MG, Araneae), by A. Brescovit & I. Cizauskas (Arachnology, 2018); 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.
:: OTHER REJECTED SPECIES: Pseudonannolene canastra (Diplopoda), synonimized under P. ambuatinga (Iniesta, Bouzan & Brescovit, European Journal of Taxonomy, 2023).
:: SPECIES WITHOUT CONFIRMATION: troglobitism not confirmed for Whiteheadiana thaisae (Carabidae, Benumea et al., Zootaxa, 2023).
:: TROGLOBITICS AWAITING ANALYSIS: Pseudochthonius limettioides (SEE).
:: INACESSIBLE INFORMATION: for Geogarypus gollumi (Geogarypidae) and Progarypus smaugi (Hesperolpiidae) in Bedoya-Roqueme et al. (Zootaxa, 2022), both Pseudoscorpiona.
:: 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) 181 formal described species not mentioned in [1], almost all described later:
Porifera (1): Arinosaster patriciae (MT, Volker et al., Neotropical Biology and Conservation, 2021).
Platyhelminthes (6): Girardia arenicola, G. corumbataiensis, G. nobresis, G. paucipunctata, G. spelaea (SP, MS, BA, A.L. Morais, M.E. Bichuette, A. Chagas-Júnior et al., Zoologischer Anzeiger, 2021); Sluysia triapertura (RN, Souza et al., Invertebrates Systematics, 2018).
Mollusca Bivalvia (1): Eupera troglobia (TO, Simone & Ferreira, Subterranean Biology, 2022).
Mollusca Gastropoda (4): Habeastrum parafusum, H. omphalium (MS, Simone, Zootaxa, 2019), Gastrocopta sharae (GO, Simone et al., Zoosystematic Evolution, 2017), Spiripockia umbraticola (BA, Simone & Salvador, Journal of Natural History, 2021).
Amphipoda (1): Spelaeogammarus ginae (BA, Bueno & Penoni, Nauplius, 2022).
Isopoda (45): Amazoniscus spica, Alboscia jotajota and Metaprosekia igatuensis (PA, SP, BA, Campos-Filho I.S et al., European Journal of Taxonomy, 2020); five Xangoniscus (BA, MG, G. M. Cardoso, Zootaxa, 2020); X. lapaensis, X. loboi (BA, Campos-Filho et al., Subterraneam Biology, 2022); Chaimowiczia tatus, C. uai (BA, MG, G.M. Cardoso (Subterranean Biology, 2021); C. obybytyra (BA, Cardoso et al., Subterranean Biology, 2022); Iansaoniscus leilae, I. paulae (BA, Cardoso et al, Subterranean Biology, 2022); six Pectenoniscus (BA, MG, G.M. Cardoso, Nauplius, 2020; Pectenoniscus lilieae, Benthana xiquinhoi (BA, Campos-Filho via et al (Zootaxa, 2019); Pectenoniscus fervens, P. pankaru (Ivanklin Soares Campos-Filho et al., ZooKeys, 2022); Spelunconiscus septemlacuum (Bastos-Pereira et al., Nauplius, 2022), Cylindroniscus platoi (Fernandes et al., Zootaxa, 2022), Gabunillo aridicola (CE/RN, Souza et al., Zootaxa, 2010), Benthana alba, Benthanoides amazonicus, B. tarzan (PA, 2023), Pectenoniscus monsviridis, P. revelatus, P. archaeos, P. sepultus, P. caesarius (Cardoso & Ferreira, Studies on Neotropical Fauna and Environment, 2023), Caraiboscia jabutiensis, Gabunillo enfurnado, Venezillo moreirai, V. limai (Campos-Filho I. S. et al, Zoosystema, 2023), Kadiweuoniscus rebellis (MS, 2024), Circoniscus mendesi, C. xikrin (PA, 2024), Ctenorillo iuiuensis (BA, 2024), Trichorhina baiana (BA, 2023).
Collembola (24): Arrhopalites glabrofasciatus, P. guanhaensis (MG, Zeppelini et al., Zootaxa, 2018), Pararrhopalites ubiquum (MG, Zeppelini, Neotrop. Entomol., 2018); 10 spp. Trogolaphysa (PA, MG, SP, Zeppelini et al. PRE-PRINT, 2021); 9 spp. of Pseudosinella (Cipola et al., Insects, 2020; see also Zeppelini et al. PRE-PRINT, 2021); Troglobentosminthurus luridus (BA, Bellini, Medeiros & Souza, Insects, 2022); Pararrhopalites sideroicus (Douglas & Brito, Florida Entomologist, 2014).
Insecta (22):
Endecous infernalis (BA, Mendes Carvalho et al., Zootaxa, 2023); Erebonyx catacumbae (BA, Melo & Ferreira, Zootaxa, 2021); Endecous troglobia (MG, Castro-Souza et al., Zootaxa, 2020), Endecous vitreus (MS, 2023).Coarazuphium auleri (MT, Pellegrini et al., Studies on Neotropical Fauna and Environment, 2021); Coarazuphium bambui (BA, Pellegrini et al., Zootaxa, 2022); Coarazuphium lundi (MG, T.G. Pellegrini et al, Zootaxa, 2020); C. xingu, C. xikrin, C. kayapo (PA, T.G. Pellegrini et al., Subterranean Biology, 2022); Perigona spelunca (MG, T.G. Pellegrini et al. (EJT, 2022); Ardistomis ferreirai (PA, Balkenohl et al., Zootaxa, 2018), Metopioxys carajas (PA, Asenjo et al., Zootaxa, 2019), Oxarthrius aurora, O. inexpectatus (Asenjo et al., Zootaxa, 2018).Mesodiplatys falcifer (Kamimura, ZooKeys, 2018).Spelaeometra gruta, Cephalometra pallida (Polhemus & Ferreira, Tijdschrift voor Entomologie, 2018), Ferricixius michaeli, F. goliathi (Ferreira et al., Zootaxa, 2023), Spelaeometra hypogea (Cordeiro, Bichuette & Moreira, Animals, 2023).
Chilopoda (2): Schendylops janelao (MG, Abreu Nunes (Zootaxa, 2019), Plutogeophilus jurupariquibaba (SP, Bonato e Ferreira, Organisms Diversity & Evolution, 2023).
Diplopoda (5): Dobrodesmus mirabilis, (BA, 2016), Phaneromerium troglopterygotum (BA, Golavatch, Gallo & Bichuette, Zootaxa, 2022), Strongylosomides troglobius (BA, Golovatch, Bouzan & Gallo, Zootaxa, 2022), Onciurosoma troglobium (Golovatch et al., Arthropoda Selecta, 2022), Cayenniola albaserrata (BA, 2024).
Acari/Trombidiforme (1): Leptus sidorchukae (MG, Santos Costa et al., Zootaxa, 2019).
Psedoscorpiones (17): Spelaeobochicha goliath (MG, Vieira et al., Zootaxa, 2018), S. mahnerti (MG, Viana & Ferreira, 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, Schimonsky, Arachnology, 2022; complementary data: Mahnert, Revue Suisse de Zooologie, 2001), Pseudochthonius koinopoliteia, P. diamachi, P. pali (Prado & Ferreira, Zootaxa, 2023), Pseudochthonius lubueno (Assis et al., Zoologia, 2023).
Palpigrades (12): Allokoenenia canhembora, A. stygia (BA, PA, Souza M.F.V.R & Ferreira R.L, European Journal of Taxonomy, 2022); Eukoenenia eywa, E. navi, E. neyriti (MG, Souza & Ferreira, Invertebrate Systematics, 2018), E. jequitai, E. lundi, E. magna (MG, Souza & Ferreira (Zootaxa, 2020), E. audax (GO, Souza, Mayoral & Ferreira, Zootaxa, 2020), E. igrejinha (MG, Souza & Ferreira, Journal of Arachnology, 2019), E. ibitipoca (MG, Souza & Ferreira, Zootaxa, 2019); Eukoenenia mocororo (MG, Souza & Ferreira, Zootaxa, 2022).
Amblypigy (5): Charinus cearensis, C. diamantinus, C. euclidesi, C. puri, C. renneri (Miranda et al., EJT, 2021).
Opilonida (4): Iandumoema cuca, I. gollum, I. stygi (MG, 2020), Spaeleoleptes gimli (BA, 2024).
Araneae (29): Ctenus igatu (BA, Cizauskas et al., Journal of Aracnology, 2022); Paracymbiomma caecus (PA, Rodrigues et al., Zootaxa, 2018); Ochyrocera ritxoco, O. ritxoo (PA, Brescovit et al., Zookeys, 2021); six Matta (MG, Brescovit & Cizauskas, Zootaxa, 2019); Carajas paraua, Tisentnops mineiro, T. onix (PA, MG, Brescovit & Ruiz, ZooKeys, 2016); Loxoscelis troglobia (BA, Souza & Ferreira, Zootaxa, 2018), Tonton itabirito, T. matodentro (MG, Pessanha et al., ZooKeys, 2013); Ochyrocera dorinha, O. rosinha, O. ungoliant (MG, PA, 2021); Speocera babau, S. pinima, S. piquira (PA, A. D. Brescovit et al., Zootaxa, 2022); Brasilomma enigmatica (MG, Brescovit et al., Zootaxa, 2012); Indiani gaspar (MG, Rodrigues et al., Zootaxa, 2020); Paracymbiomma bocaina (PA, Rodrigues et al., Zootaxa, 2018), Paracymbiomma pseudocaesus (PA, Cizauskas, Dissertation, 2017), Loxosceles bodoquena (MS, 2024), Pinelema elinae (BA, 2024), Loxosceles boqueirao (BA, 2024).
Pisces (1): Phreatobius sanguijuela (Zampulo & Prous (Fauna Cavernicola do Brasil, 2022).
B) 327 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.
MG
Cordisburgo 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).
TO
Tocantins (6 spp., Dianópolis, [TO/1], Ferreira et al., CECAV, 2016).
MS
Mato Grosso do Sul (31 spp., [MS/1], Cordeiro et al., Biota Neotropica, 2014).
RN
Rio Grande do Norte state (51 spp., [RN/1], Bento et al, Biodiversity and Conservation, 2022; EXCEL LIST); Potiberaba porakuara also in Ceará).
ES
Espírito Santo state (10 spp., [ES/1], SEE).
GO
São 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).
SP
Alto do Ribeira region in SP (six Potamolithus (Lithoglyphidae), [SP/1], Maria Elina Bichuette & Eleonora Trajano, Subterranean Biology, 2018).
MIXED
Campo 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 troglobite 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.
Some inventories result in confirmation of the absence of troglobites 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 troglobitic/trogomorphic species in Minas Gerais state, however, without discriminating which are exact troglobites, we decided not to count these taxa. Trevelin et al. (Ecological Indicators, 2019) quotes an incredible 135 possible troglobites in Serra dos Carajás (PA), among Amblypygy (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 troglobites, bringing Brazilian diversity to c. 720 spp. However, we do not include this record here due to the lack of a more detailed inventory.
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 (46) ‣ BA (16), MG (13), RN (9, 1 non-endemic), SP (2), ES (2), GO (2), MS (1), TO (1), CE (1, non-endemic).
Collembola (46) ‣ MG (16), BA (9), RN (9), MS (5), SP (4), GO (3).
Diplura (9) ‣ MG (3), RN (3), BA (3).
Insecta (41):
Zygentoma ‣ BA (3), ES (2), MG (1).
Coleoptera ‣ MG (10), BA (6), RN (2), MS (1), SP (1), GO (1), TO (1).
Blattodea ‣ BA (3), MG (2).
Hemiptera ‣ MS (1), BA (3).
Hymenoptera ‣ BA (1), TO (1).
Pauropoda (1) ‣ MG (1).
Symphyla (2) ‣ SP (1), MG (1).
Diplopoda (52) ‣ BA (15), 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 (30) ‣ BA (11), 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 (6) ‣ BA (4), MG (1), SP (1).
Opiliona (16) ‣ MS (6), MG (4), BA (3), ES (2), RN (1).
Pisces (5) ‣ MS (3), BA (2).
BY GROUP
Largest groups, including non state-endemic species: Isopoda (103), Collembola (84), Diplopoda (65), Araneae (61), Coleoptera (46), Pseudoscorpiona (39), Opiliones (30), Pisces (27), Palpigradi (24), Chilopoda (21), Amphipoda (21), Gastropoda (21), Platyhelminthes (13), Hemiptera (12), Diplura (10).
BY STATE
Only 14 troglobites are non states-endemic: 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 (196), Minas Gerais (180), São Paulo (64), 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).
If we count all the species (inc. non state endemics), we have, the numbers are Bahia (197), Minas Gerais (184), São Paulo (71), 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 (103), Minas Gerais (84), Rio Grande do Norte (51, one non-endemic), São Paulo (32), Mato Grosso do Sul (31), Espírito Santo (10), Goiás (10), 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]. 148 municipalities has troglobitics 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 ‣ 25 municialities: Carinhanha (54), Andaraí (38), Iuiu (30), Campo Formoso (26), Malhada (11), Coribe (9), Iraquara (6), Itaetê (6), Santana (5), São Desidério (5), Feira da Mata (4), Serra do Ramalho (4), Morro do Chapéu (3), Palmeiras (3), Paripiranga (3), Lençóis (2), Varzea Nova (3), Bom Jesus da Lapa (1), Curação (1), Ituaçu (1), Miranguara (1), Nova Redenção (1), Ourolândia (1), Pau Brasil (1), Santa Maria da Vitória (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), Arinos (7), Mariana (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), Luislandia (4), Caete (3), Curvelo (3), Lagoa Santa (3), Santa Bárbara (3), Ibiracatu (2), Itambé do Mato Dentro (2), Monjolos (2), Prados (2), Prudente de Morais (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 (51), 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 (54), Iporanga (51), Felipe Guerra (41), Cordisburgo (39), Andaraí (38), Iuiu (30), Campo Formoso (26), Bonito (24), Bodoquena (23), Parauapebas (21), Pains (17), São Domingos (17), Itacarambi (16), Canaã dos Carajás (13), Itu (12), Malhada (11), Matozinhos (11), Rio Acima (11), Belo Horizonte (10), Coribe (9), Baraúna (9), Governador Dix-Sept Rosado (9), Itabirito (9), Arinos (7), Mariana (7), Pedro Leopoldo (7), Barão de Cocais (6), Brumadinho (6), Curionópolis (6), Dianópolis (6), Iraquara (6), Itaetê (6), Morro do Pilar (6), Nova Lima (6), Sete Lagoas (6), Vazante (6), São Desidério (5), Conceição do Mato Dentro (5), Jardins (5), Lima Duarte (5), Montes Claros (5), Presidente Olegário (5), Santana (5), Vargem Alta (5), Feira da Mata (4), Luislandia (4), Porto Murtinho (4), Serra do Ramalho (4), Caete (3), Caete (3), Corumbá (3), Curvelo (3), Eldorado (3), Lagoa Santa (3), Mambaí (3), Morro do Chapéu (3), Palmeiras (3), Paripiranga (3), Posse (3), Rio Branco do Sul (3), Santa Bárbara (3), São Felix do Xingu (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), Prudente de Morais (2), Quixerê (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), Cantagalo (1), Cambuci (1), Conceição do Castelo (1), Capim Branco (1), Caraí (1), Catas Altas (1), Coronel José Dias (1), Curação (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), Miranguara (1), Moeda (1), Montalvania (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 da Vitória (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), Varzea Nova (1). Vespasiano (1).
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).
 |
| BRAZILIAN TROGLOBITIC LOCATIONS (INC. BOLIVIAN RECORD OF PHREATOBIUS SANGUIJUELA). DUE TO SCALE, EACH POINT MAYBE SEVERAL CAVES/MUNICIPALITIES. |
Important note: no photos used on this page of troglobionts 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.
5.1 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 (a new genus at their description, Volker et al., Neotropical Biology and Conservation, 2021); and Racekiela cavernicola Volkmer-Ribeiro, Bichuette & Machado, collectd in Lapa dos Brejões, northern Bahia state (Volker et al., Neotropical Biology and Conservation, 2010); both troglobite species cited here are corroborated by [6]; a fourth species, undescribed, was discovery in center Mexico (Legendre et al., Subterranean Biology, 2023).
5.2 PHYLUM CNIDARIA ‣ many records of Cnidaria in freshwater caves can be found in Zagmaster et al. (Speleobiology, 2011), in several countries: Mexico, U.S.A., Australia and several center European countries; the hydra Velkovrhia enigmatica Matjasic & Sket, 1971 (Bougainvilliidae, PHOTO) is the only freshwater hydrozoan living exclusively in groundwater; it is endemic to the Dinarides in the Balkan Peninsula, where it has been known from five caves in Slovenia (3), Croatia (1) and Bosnia (1), by Magmajster, Natura Sloveniae, 2003; strangely, it is not listed in [3] for freshwater Cnidaria.
5.3 PHYLUM PLATYHELMINTHES ‣ near c. 165 spp. of flatworms are troglobitics, c. 150 in Tricladida, c. 15 in Temnocephalida (parasites on Troglocaris in S Europe and one at a crab in Papua New Guinea, [2, pg. 1176]), and one Prorhynchidae in South Africa (Ferreira et al., Subterranean Biology, 2020).
All (3:3/)13 Brazilian obligatory subterranean Platyhelminthes belongs order Tricladida [1] in all three clades: Continenticola (10 in Girardia, Dugesiidae), Maricola (1) and Cavernicola (2). Sluysia triapertura Leal-Zanchet & Souza, 2018 represents the first Maricola triclad living in freshwater within a cave (Souza et al., Invertebrates Systematics, 2018). Brazil is the only country in the world with troglobitic representatives in the three groups of Tricladida. Four undescribed spp. was cited from Mato Grosso do Sul [MS/1], São Paulo [SP/2], these Continenticola, one Cavernicola for Rio Grande do Norte [RN/1], and one of undetermined order for Minas Gerais [MG/7]. Paraba pankaru Amaral & Leal-Zanchet, 2019 (Continenticola, S. V. do Amaral et al., SEE) is a troglophilic.
(5:6/)33 cave spp. occur in North America, mainly in genus Spalloplana; cave species absents in China. Six spp. occur in Mexico, all Tricladida (Mexican Cave Fauna, 2022).
UNDETERMINED TURBELLARIA
CAVERNICOLA
CONTINENTICOLA
MARICOLA
5.4 PHYLUM NEMATODA ‣ 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 troglobiont. Carefully consulting the Du Preez (Nematology, 2017), only seven spp. possibly are true troglobitics: 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), Chronogaster troglodytes Poinar & Sarbu, 1994 (Romania). Axonchium sbordonii, from Sima del Ojito, Chiapas, Mexico, is rejected in this work (Mexican Cave Fauna, 2022). In adition, one possibly troglobitic species was collected in Colombia (Angarita-Sierra, CCE/2018).
5.5 PHYLUM NEMERTEA ‣ by [2], 3 of all 20 freshwater Nemertea are possibly troglobitics (stygobionts): Prostoma puteale in France, Switzerland, and Germany, and Prostoma hercegovinense in Bosnia and Herzegovina, but one species has been described from New Zealand also. In addition, Brazil has two possibly troglobitic 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 troglobite Nemertea described for the New World.
5.6 PHYLUM ANELLIDA ‣ some polychaete families have adapted to living in the water column of marine/anchialine caves, including the normally benthic Protodrilidae, Nerillidae, and Scalibregmatidae. Marifugia cavatica Absolon & Hrabe, 1930 (Serpulidae) is eyeless and lacks pigmentation, and is also considered to be a Tertiary relict of marine origin. The aptly named genus Troglochaetus (Nerillidae) is also found in the hyporheic zone of caves, wells and springs, with T. beranecki 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). 17 cave spp. in 6 genera of 3 families occcur in in North America [2].
A. Sabellida has Marifugia cavatica, known from caves situated in Dinaric Karst (northeastern Italy, Slovenia, Croatia, Bosnia, and Hercegovina) but occurs in small, isolated areas, only known freshwater and stygobiotic member of group.
B. Phyllodocida has at least 12 stygobiotic (however, all inconcluse be troglobionts) species representing genus Namanereis. C. J. Glasby et al. (ZJLS, 2014) lists all stygobiont Nemanereis: N. cavernicola 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 in Oman, two in Socotra, N. pilbarensis in W. Australia, N. berone in Papua New Guinea and N. tiriteae in New Zealand and Fiji.
Clitellata ‣ aquatic clitellates 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, the stygobiotic (i.e., obligatory hypogean) nature of the species can only be inferred from their exclusive presence in the subterranean environment. Despite their morphological pre-adaptation, only approximately 100 spp of aquatic oligochaetes are exclusively present in groundwater out of approximately 1,700 aquatic oligochaete species and 1100 freshwater species known to date. In U.S.A. occur 11 spp. in 5 genera among 2 families. A checklist of stygobiotic Oligochaeta from caves is disponible in Creuzé des Chatelliers (Freshw. Biol., 2009), with 112 spp. in 7 families. In New World are 20 spp., in four families. Possibly 4 troglobitic Clitellata occur in Brazil, none formally described, two in Bodoquena region in Mato Grosso do Sul state [MS/1], and two in W Rio Grande do Norte state [RN/1]. No data from terrestrial troglobitic Oligochaeta; at least two occur in Mexico (Mexican Cave Fauna, 2022).
C. Clietellata/Tubificida: two families with cave species: Phreodrilidae ((3/)5 spp., in New Zealand, Oman, Marocco and Australia) and Tubificidae ((19/)49 spp., 10 spp. in New World: Dero haitiensis, Spirospermoides stocki (Haiti and adjacent islands), four Pristina, Clitellio cavernicolus (Cuba), Gianius aquaedulcis (France, Germany, Italy, Spain, U.S.A./New York), Phallodriloides macmasterae (Bermuda), Krenedrilus papillatus (Venezuela).
D. Clietellata/Haplotaxida: a single family with cave species: Haplotaxidae ((3/)8 spp., in Europe, Japan and Guinea).
E. Clietellata/Parvidrillida: one family with cave species: Parvidrilidae (two spp. in Parvidrilus, one in Italy another in Alabama, U.S.A.).
F. Clitellata/Enchytraeidae: (6/)13 spp.in caves, all in Europe except Marionina ecuadoriensis Righi and Hemienchytraeus mauriliae Righi from Ecuador.
G. Clitellata/Lumbriculidae (inc. Dorydrillidae): (9/)38 spp., mainly in Europe, one in Japan, 7 in U.S.A., in Virginia, Alabama Tennessee, Oregon and California: Spelaedrilus multiporus Cook, Stylodrilus beattiei Cook, S. californianus Rodriguez, S. wahkeenensis Rodriguez & Coates, Tenagodrilus musculus Eckroth & Brinkhurst, Trichodrilus allegheniensis Cook and T. culveri Cook.
H. Clitellata/Hirudina/Hirundinina: troglobionts in this group usually exhibit two specific morphological features, namely milky-white coloration and lack of eyes or eye pigment (Cichocka et al., Zootaxa, 2015); troglobitic leeches cited in literature fall in this description: Haemopis caeca from Dobrogea in Romania, some Erpobdellidae present in southern Europe (N Italy–Balkans–Turkey–Caucasus, possibly Erpobdella absoloni Johansson, 1913), E. borisi Cichocka & Bielecki from NW Iran, E. krasensis Sket, 1968 and E. mestrovi Kerovec, Kučinić & Jalžić, 1999 from Croatia (Cichocka, 2015), with possibly undescribed species, probably erpobdellids, in caves of China and the U.S.A. (Sket & Trontelj, Hydrobiologia, 2006); however, 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; the species inhabit the aphotic zone of a caves in Papua New Guinea and China, respectively (Cichocka, 2015). Two species of two diferent families are troglobites in China. Two possibly undescribed troglobitic species was collected in C Minas Gerais state [MG/1] and [MG/7], Brazil; if confirmed, they will be the firsts troglobitics Hirudinea described for the New World.
I. Clitellata/Hirudina/Branchiobdellida (crayfishworms) are also known from caves as stygobiotic; 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 two spp. occur in Mexico (Mexican Cave Fauna, 2022) and six in U.S.A. [2].
5.7 MOLLUSCA/BIVALVIA ‣ typical troglobitic mollusk 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 troglobitic species. Bivalvia includes only two troglobitics described: 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 troglobitic bivalvia in western Hemisphere.
5.8 MOLLUSCA/GASTROPODA ‣ all terrestrial troglobite snails are all pulmonates, except for the prosobranch Pholeoteras euthrix, has so far been known from caves in southern Croatia, southern Herzegovina and Greece (ANIMALBASE). The number of terrestrial cave-limited species is much less than the aquatics, probably less than 50, with hotspot for terrestrial cave snail fauna is in SE Asia; there are over 450 described species of stygobitic snails, almost all prosobranchs.
In the Slovenian there are 37 aquatic obligate cave snails, and 11 of terrestrial obligate cave snails; France which is home to 143 stygobite snails, most of them described in the last 40 years. In U.S.A. occur (5:16/)32 sp. of obligate-cave gastropods, 27 stygobionts and 5 terrestrial [2]; in Mexico, all troglobitic (3:5/)9 gastropoda are stygobionts (Mexican Cave Fauna, 2022); China has two troglobitic Mollusca, both Pulmonata, in two families [2]. Recent data point to (5:20/)39 spp. for USA/Mexico, by Gladstone et al. (Conservation Biology, 2022).
Cavallaria et al. (Zoologia, 2022) cites 19 described troglobitic gastropoda from Brazil, however, troglobiontism for many of them are rejected by several autors. Here we accept 21 names as troglobionts in country, 6 already described and mentioned expressly in R.L.Ferreira et al. (Zoologia, 2023), and 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]. Overall, Brazil has at least 8 genera and 8 families.
UNDETERMINED FAMILIES
CAENOGASTROPODA
DIPLOMMATINIDAE
EUPULMONATA
CHAROPIDAE
COCHLIOPIDAE
GASTROCOPTIDAE
LITHOGLYPHIDAE
POMATIOPSIDAE
SCOLODONTIDAE
SYSTROPHIIDAE
5.9 PHYLUM ONYCHOPHORA ‣ by [4], officially only two trogobitic Onychophora are known: Peripatopsis alba (Peripatopsidae) of South Africa and Speleoperipatus spelaeus (Peripatidae) from Jamaica, with probable occurrences in the Galapagos; unofficially, there are probably troglobitic populations also in Brazil, in caves of the Bodoquena System, in Mato Grosso do Sul, of a species that has not yet been formally described, and may even be a new genus [MS/1].
PERIPATIDAE
5.10 ARTHROPODA/ARACHNIDA ‣ all orders of Archnida has troglobites except Holothrida and Solifugae. 337 troglobitic 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 troglobitic species: Parasitiformes with the orders Opilioacarida and Gamasida, and Acariformes with the orders Sarcoptiformes and Trombidiformes. U.S.A. has (10:17/)26 spp. troglobitic 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 troglobitic 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 troglobitic, despites several cave collections (Berbardi & Borges-Filho, Subterranean Biology, 2018); cave members occur only in Cuba, Thailand and Vietnam [4].
B. Ixodidde ‣ by Galán & Herrera (BSVE, 2006) one species (Antricola silvae) from Venezuela maybe a troglobitic. If confirmed, it will be the only record of an Ixodida troglobite worldwide. Huge populations occur in caves in Brazil, associated with bats.
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 non described species in Brazil, collected in Minas Gerais [MG/1] and Bahia [BA/3] states.
DITHINOZERCONIDAE
PACHYLAEPIDAE
D. Trombidiformes ‣ Prostigmata (more than 20,000 species), 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 troglobite, Proterorhagia oztotloica from Mexico; troglobitic Rhagidiidae are often linked to cold caves in the Holarctic [4]. One troglobitic 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 troglobitic Hydracnidia in three families (Galán & Herrera, BSVE, 2006).
ERYTHRAEIDAE
RHAGIDIIDAE
LABIDOSTOMATIDAE
E. Sarcoptiformes ‣ cave-restricted species of this group are surprisingly rare: some species of Schwiebea (Acaridida) from humid hypogean habitats, devoid of adaptations to subterranean life, and a few European Belbidae, such as Metabelbella phalangioides - pale and long-legged [4]. Brazil has two undescribes species in Mato Grosso do Sul [MS/1] and Bahia [BA/3].
HIDROZETIDAE
OEHSERCHESTIDAE
F. Pseudoscorpiones ‣ c. 400 troglobitic species in c. 15 families was described. New World troglobitic falls in 12 of them. Mexico has 40 spp. in 5 families; U.S.A. has 150 spp. in 29 genera of 9 families; (4:)39 spp. in Brazil [2], 15 undescribeds, in Bahia [BA/2][BA/3][BA/4], Minas Gerais [MG/1][MG/6][MG/10], Rio Grande do Norte [RN/1], São Paulo [SP/3] and Goiás [GO/1].
BOCHICHIDAE
CHTHONIIDAE
CHERNETIDAE
IDEORONCIDAE
SYARINIDAE
G. Araneae ‣ c. 1,000 cave species in 48 families, 194 in Europe [4]. U.S.A. has (10:22/)119 spp.; Cicurina (Dictynidae) has 55 spp. troglobitics in Texas [2]. Mexico has (11:)59 spp.; Colombia includes 3 spiders (in 3 families, Pisauridae without Brazilian troglobitics, Angarita-Sierra, CCE/2018, also Angarita-Sierra et al, Chapter Book, 2019).
Brazil has (16:)64 spp., 30 undescribeds in country, in Bahia [BA/1][BA/2][BA/3][BA/4][BA/5], Minas Gerais [MG/1][MG/3][MG/5][MG/10], Rio Grande do Norte [RN/1], Mato Grosso do Sul [MS/1], São Paulo [SP/2], Tocantins [TO/1], and Espírito Santo [ES/1].
Among Theraphosidae, troglobitc species includes only six Hemirrhagus from Mexico and one Tmesiphantes from Brazil (Bertane, Bichuette & Pedroso, Anais da Acadêmia Brasileira de Ciências, 2013). Besides several collections inside caves, seven spp. of Ochyrocera describeds in Brescovit et al. (Zootaxa, 2018) and three Speocera describeds in A.D. Brescovit et al. (Annales de la Société entomologique de France, 2023) are not troglobitic.
UNDETERMINED FAMILY
CAPONIIDAE
CTENIIDAE
DIPLURIDAE
HAHNIIDAE
MICROSTIGMATIDAE
NESTICIDAE
OCHYROCERATIDAE
OONOPIDAE
PALPIMANIDAE
PHOLCIDAE
PRODIDIMIDAE (syn. GNAPHOSIDAE)
SICARIIDAE
SYMPHYTOGNATHIDAE
TELEMIDAE
TETRABLEMMIDAE
THERAPHOSIDAE
H. Opiliones ‣ in order Cyphophthalmi the unique New World member from caves is Neogovea mexasca from Mexico; Eupnoi no has true troglobites; Dyspnoi (Palpatores) has only two possible troglobites species in New World: one Nemastoma in U.S.A. and Ortholasma sbordonii in Mexico; all remaining troglobitc opiliones in New World belongs Laniatores [2.4].
Mexico has (4:)16 spp. [4]. U.S.A. has (7:13/)39 spp. [2]. Angarita-Sierra (CCE/2018) and Angarita-Sierra et al (Chapter Book, 2019) cites four troglobitic Opiliona (in four families) for Colombia; Venezuela includes 4 spp., all in Trinella at Agoristenidae (Galán & Herrera, BSVE, 2006). Brazil has (8:)31 spp., all Laniatores, 16 undescribeds, in Bahia [BA/2][BA/3], Minas Gerais [MG/1][MG/3][MG/6], Espírito Santo [ES/1], Mato Grosso do Sul [MS/1] and Rio Grande do Norte [RN/1].
COSMETIDAE
CRYPTOGEOBIIDAE
ESCABIIDAE
GERDESIIDAE
GONYLEPTIDAE
KUMULIDAE
TRICOMMATIDAE
ZALMOXIDAE
I. Amblypigy ‣ troglobites are cited in four families, three in New World: Charinidae (Charinus, inc. all 11 Brazilian species; absents in Mexico; two spp. in Venezuela, by Galán & Herrera, BSVE, 2006); Paracharontidae, with Jorottui, troglobitic endemic to N Colombia (Novataxa, 2023), and Phrynidae, with true troglobitics in Mexico (11), Cuba [4] and Colombia (two, Angarita-Sierra, CCE/2018). No troglobitic Amblypigy occur in U.S.A.
CHARINIDAE
J. Schizomida ‣ the two families of this order has troglobitics: Hubbardiidae has unquestioned troglobites are known only from caves in Cuba, Jamaica, Belize, Mexico (14), California (1, U.S.A.), Ecuador, and Australia; in Protoschizomidae, all 10 spp. are troglobitics, 9 in Mexico and one in U.S.A [2].
L. Thelyphonida ‣ no cave dwellers were reported in this order, but possibly one sp. occur in Laos [4].
L. Palpigradi ‣ troglobitic Palpigrada occur in Europe (22), Thailand (4), Cuba (1), Indonesia (2), India (10), and Brazil (24); all genera in this order except Leptokoenenia have troglobites [2]. In New World cave species occur only in Brazil (two in Allokoenenia, 22 in Eukoenenia) and Cuba, with possibly undescribed species in California [2] and Mexico [4]. 6 undescribeds in Brazil, in Minas Gerais [MG/1], Bahia [BA/1][BA/2][BA/3] and São Paulo [SP/2].
EUKOENENIDAE
M. Ricinulei ‣ troglobites in Pseudocellus have been described from Cuba (1), Belize (1) and Mexico (11). No troglobites occur in Brazil.
N. Scorpionida ‣ there are 24 troglobitic scorpions worldwide, in Mexico (13, [5]), Indonesia (2), Australia (2), Israel (1), Madagascar (1), U.S.A. (Uroctonus grahami, Vaejovidae), Ecuador (Troglotayosicus vachoni, Troglotayosicidae), Venezuela (Taurepania trezzii, Chactidae, Galán & Herrera, BSVE, 2006), Colombia (Tityus grottoedensis, Angarita-Sierra, CCE/2018) and Brazil (Troglorhopalurus translucidus, Buthidae). An additional six species known only from caves show slight reduction in pigmentation or eyes but are probably not troglobitic [2].
BUTHIDAE
5.11 ARTHRPODA/PAUROPODA ‣ no known species of obligatory subterranean [2], but one species may be a troglobitic; one undescribed species was collected in Minas Gerais [MG/1], Brazil; if confirmed, it will be the first Pauropoda troglobite described by science.
5.12 ARTHROPODA/SYMPHILA ‣ only three species of this order are described as possibly obligatory subterranean, one in Tasmania (Hanseniella magna, SEE) and two in Slovenia; two undescribed collected species from Brazil in Alto do Ribeira karst, São Paulo state [SP/3], and C Minas Gerais [MG/6] are possibly triglobites - if confirmed, it will be the first Symphyla troglobite described in New World.
5.13 ARTHROPODA/DIPLOPODA ‣ no conclusive data on the world diversity of troglobiont Diplopoda was found by Brazilian Metazoa, but it is known that it affects several hundred species. 11 of the 16 orders of the class has troglobitics. In Brazil four orders have obligatory subterranean species.
(13:32/)133 troglobitic Diplopoda are described in North America in 5 orders [2], 114 in China [2], (15:)73 in Mexico [5, in six orders], 4 in Colombia (in four distinct families, Angarita-Sierra, CCE/2018) and (at least 14:)65 in Brazil in 5 orders, 52 undescribeds.
Heterocaucaseuma deprofundum Antic & Reboleira is a recently discovered troglobiont from deep caves in the Caucasus, and at 1,980m below the surface is the deepest occurring known millipede and perhaps the deepest occurring terrestrial arthropod.
UNDETERMINATE ORDERS
POLYXENIDA ‣ abundant in dry soils and under barks, with ca. three troglobites: Lophoproctus pagesi from Balearic Islands and two recently described species from Christmas Island (Lophoturus speophilus and L. humphreysi) [2]; two undescribed species was collected in Bahia [BA/4] and Minas Gerais [MG/7].
POLYXENIDAE
HYPOXEGENIDAE
GLOMERIDA ‣ troglobites in this order are common troglobionts in Europe, China, Mexico, and Central America. 26 troglobites are collected in Old World; in New World only the genus Glomeroides is present underground, with six microphthalmic troglobites from Mexico [2, 4].
SIPHONOPHORIDA ‣ four caves species: one in Mexico, two in California, U.S.A., these they are phyletically eyeless, but not troglomorphic [2, 4]; and one in Brazil, possibly troglobitic, undescribed in Bahia [BA/5].
SIPHONOPHORIDAE
GLOMERIDESMIDA ‣ a smaller order, with only two troglobitc species, Glomeridesmus sbordonii Shear, 1974 from Mexico and G. spelaeus Iniesta & Wesener, 2012, from Curionópolis, SE Pará state, in N Brazil (Iniesta, Zootaxa, 2012) [2,4].
GLOMERIDESMIDAE
JULIDA ‣ dominant millipede order in Europe and many troglobionts have been described; there are at least two troglobiont species in North America as well, but relatively few in Asia, and none are known from Mexico [2,4].
SPIROBOLIDA ‣ this order has has at least two troglobites: the troglomorphic guanobite Reddellobus troglobius in Mexico and Speleostrophus nesiotes, depigmented and eyeless, in Australia [2,4].
CALLIPODIDA ‣ troglobiotic species in this order are found in North America, China, Vietnam, and the Middle East. Absents in Mexico [2,4].
SPIROSTREPTIDA ‣ this order is subdivided into Cambalidea (5:242 spp.) and Spirostreptidea (5:772 spp.). Cambalidea includes a high proportion of cave-related species (inc. 4 in Mexico), while Spirostreptidea has only three troglobites from S Mexico caves; truly troglobitics are only three: Cambala speobia Chamberlin, 1953 and C. reddeli inornatus Causey, 1964 both rom gypsite caves in Texas (USA) and Mexicambala russelli Causey, 1964 found in caves in Mexico (Iniesta & Ferreira, Zootaxa, 2013). In South America all troglobitc are Pseudonannolene from Brazil [2]; 4 undescribed species occur in caves from Goiás [GO/1], Espírito Santo [ES/1] and Bahia [BA/2].
PSEUDONANNOLENIDAE
CHORDEUMATIDA ‣ the most important millipede order in north temperate zone caves is the Chordeumatida; in the U.S.A. alone, there may be more than 100 troglobiont chordeumatidan species, about a third of them undescribed. 13 troglobitics in Mexico [2,4]
STEMMIULIDA ‣ besides some cave-dwelling species (e.g. Stemmiulus in N Brazil), possibly none troglobites belongs this order [2,4].
POLYDESMIDA ‣ distributed worldwide, Polydesmida has subterranean representatives in all temperate and tropical regions except Sub-Saharan Africa. No species seems to be guano dependent, but troglobites and troglophiles are plenty; many troglobiont species have been described from Europe, Japan, China, and Mexico, and they are the dominant invertebrate detritivores in many Mexican caves; in South America many species occur in several families, especially in south Brazil, include numerous troglobites. 45 troglobitics in Mexico in this order.
A large number of undescribed species were collected in caves in many states of Brazil: Bahia [BA/1][BA/2][BA/3][BA/4], Espírito Santo [ES/1], Goiás [GO/1], Mato Grosso do Sul [MS/1], Minas Gerais [MG/1][MG/3][MG/5][MG/6][MG/10], Rio Grande do Norte [RN/1], São Paulo [SP/2][SP/3][SP/4]. One species of Trichopolydesmidae is a troglobitic in Venezuela (Galán & Herrera, BSVE, 2006).
UNPLACED AT FAMILY LEVEL
CHELODESMIDAE
CRYPTODESMIDAE
DOBRODESMIDAE
ONISCODESMIDAE
PARADOXOSOMATIDAE
POLYDESMIDAE
PYRGODESMIDAE
SPHAERODESMIDAE
TRICHOPOLYDESMIDAE
5.14 ARTHROPODA/CHILOPODA ‣ of the five orders of Chilopoda, four have cave species, all of them in Brazil; for cave Chilopoda worldwide, see Chagas and Bichuette (Zookeys, 2018). 22 troglobitics occur in Brazil, (3:4/)4 in North America [2], (2:)5 in Mexico [5, two Lithobiidae, 3 in Newportia] and only one in China [2] and Colombia (Angarita-Sierra, CCE/2018, undescribed Scolocryptops). All six families with troglobitic has species in Brazilian caves except Balophilidae [4].
ORDER SCUTIGEROMORPHA ‣ Sphendononema sp. collected in Bahia state possibly representes the unique troglobitic Scutigeromorpha worldwide.
PSELLIOLIDAE
ORDER LITHOBIOMORPHA ‣ only two families belongs this order: Henicopidae and Lithobiidae; following [2], about 50 spp. troglobionic species of this order (all Lithobiidae) occur in many countries in Europe, and two spp. occur in the Maghreb, three in North America, and remaining from E Asia; two undescribed species was collected in Bahia [BA/2] and Rio Grande do Norte [RN/1] states in Brazil.
ORDER SCOLOPENDROMORPHA ‣ five families in this order, two of them has troglobitics: Cryptopidae: Cryptops longicornis (Spain), C. vulcanicus (Canaries, Spain), C. umbricus (Italy), C. roeplainsensis (Australia), C. camoowealensis (Australia), C cavernicolus (Cuba), C. troglobius (Cuba), C. iporangensis (São Paulo, Brazil) and C. spelaeoraptor (Bahia, Brazil); Scolocryptopidae: Newportia leptotarsis (Cuba), N. troglobia (Mexico), N. stoevi (Puerto Rico), N. spelaea (Bahia, Brazil), N. potiguar (Rio Grande do Norte, Brazil), Scolopocryptops troglocaudatus (Bahia, Brazil), Thalkethops grallathrix (U.S.A.). Two undescribed species occur in Brazil, in Bahia state [BA/2][BA/3].
CRYPTOPIDAE
SCOLOCRYPTOPIDAE
ORDER GEOPHILOMORPHA ‣ 13 families; among this group only five troglobitic species ae known, by Nunes et al., Zootaxa, 2019: Plutogeophilus jurupariquibara (Brazil), Geophilus persephones (France) and G. hadesi (Croatia) in Geophilidae; Ityphylus cavernicolus (Cuba) in Ballophilidae; and Schendylops janelao in Schendylidae, known only from nortern Minas Gerais state, Brazil. 5 undescribed species occur in Bahia [BA/1][BA/2], São Paulo [SP/3], Rio Grande do Norte [RN/1] and Minas Gerais [MG/5].
GEOPHILOMORPHA UNDETERMINED FAMILY
GEOPHILIDAE
SCHENDYLIDAE
5.15 ARTHROPODA/OSTRACODA ‣ many species are subterraneans; few informations, but possibly c. 600 are stygobitics [2, pg. 334]. (2:11/)16 cave species occur in North America [2], (3:3/)4 in Mexico [5, in two orders], none in China; one undescribed species was collected in Rio Grande do Norte [RN/1], Brazil. Venezuela has two troglobitic Ostracoda, both Cyprididae (Galán & Herrera, BSVE, 2006).
5.16 ARTHROPODA/MAXILLOPODA 1 ‣ this class includes six high variable subclasses, only three with cave species.
Mystacocarida ‣ none informations.
5.17 ARTHROPODA/MAXILLOPODA 2 ‣ this class includes six high variable subclasses, only three with cave species.
Copepoda ‣ few informations, but possibly c. 1,100 are stygobitics [2, pg. 334]; (4:7/)17 cave species occur in North America and 23 in China [2]. (5:8/)12 spp. occur in Mexico; in Brazil at least 4 undescribed troglobitic Copepoda are collected, in Rio Grande do Norte [RN/1], Minas Gerais [MG/6] and São Paulo [SP/3]. Venezuela includes one troglobitic Copepoda, in Cyclopidae (Galán & Herrera, BSVE, 2006).
UNKNOWN GROUP
HARPACTICOIDEAE
CYCLOPOIDEA/ENTOBIIDAE
Tantulocarida ‣ there is only one species of troglobitic (stygobiotic) tantulocarid (Stygotantulus stocki), which was reported from anchialine environments of the lava tubes in the Canary Is., parasitizing two families of harpacticoid copepods [2].
5.18 ARTHROPODA/MALACOSTRACA ‣ 13 of 17 orders has obligate-cave species; Brazil and China has obligate subterranean in Amphipoda, Isopoda and Decapoda; Brazil also has in Spelaeogriphacea and Bathynelllacea, and China in Bathynellacea [1][2]. 346 obligate subterranean species in North America, 68 in China [2], 104 in Mexico [5] and 121 in Brazil.
Leptostraca, Bochusacea, Cumacea, Myctacea and Tanaidacea has cave spcies only for marive cave systems, mainly in Caribbean region. Amphipoda, Isopoda and Decapoda occur widely in over countries with high amount of troglobitics (inc. Brazil, China, U.S.A., Mexico). Bathynellaceae, Thermosbaenacea and Mysida are relatively widely in some regions. Anaspidacea and Spelaeogriphacea are very rare.
1. LEPTOSTRACA ‣ a single troglobitic (stygobitic) species in this order is known, from Providenciales caves in the Turks y Caicos.
2. BATHYNELLACEAE ‣ 80-90 spp. are subterranean. U.S.A. has (2:8/)19 stygobitic species, and China has one cave species [2]; troglobitic Bathynellaceans absents in Mexico. Brazil has a undescribed species collected in Mato Grosso do Sul state [MS/1]. One troglobitic Parabathynellidae was collected in Venezuela (Galán & Herrera, BSVE, 2006).
3. ANASPIDACEAE ‣ confined to Oceania, few species are cave dwelling in Australia and Tasmania.
4. AMPHIPODA ‣ worldwide are c. 900 subterranean species, China has (5/)26 spp. and U.S.A. has (9:19/)182 spp. [2]; from Brazilian caves there are currently (4:5/)21 spp. of this order: Hyalella (5), Megagidiella (1), Potiberaba (5), Spelaeogammarus (8) and Seborgia (1). Mexico has (2:)22 troglobitic species in Bogidiellidae and Hadziidae. Undescribed species was collected in Rio Grande do Norte state [RN/1]. Venezuela includes three troglobitic Amphipoda in three families: Bogidiellidae, Hadziidae (Metaniphargus venezuelanus, anchialine, the only representative of Hadziidae in the underground waters of South America) and Hyalellidae (Galán & Herrera, BSVE, 2006).
AMPHIPODA/ARTESIIDAE
AMPHIPODA/HYALELLIDAE ‣ family with 12 genera. Hyalella has 70 spp., all from New World, 22 in Brazil. Currently, there are only seven species of the genus inhabiting exclusively caves: H. anophtalma Ruffo, in NW Venezuela, H. muerta Baldinger, Shepard & Threloff from California, and five species in Brazil (Rodrigues et al., Zootaxa, 2014).
AMPHIPODA/MESOGAMMARIDADE ‣ a small family known only in Korea, Japan, Indian subcontinent, southern Alaska and caves from Rio Grande do Norte state in NE Brazil (C. Fiser et al. (Systematics and Biodiversity, 2013); for recent notes: Rocha (Dissertation, 2021).
AMPHIPODA/SEBORGIDAE ‣ another small family known only in Vietnan, islands in adjacences of Papua New Guinea, Texas and caves from from Rio Grande do Norte state in NE Brazil (C. Fiser et al. (Systematics and Biodiversity, 2013).
5. THERMOSBAENACEA ‣ organisms are known from anchialine environments, caves, cenotes, interstitial environments; and springs of thermal or cold waters; Mexico and U.S.A. has a single Thermosbaena each, both confined to caves: Tulumella unidens from Quintana Roo, and Monodella texana from Texas, respectivey [2].
6. SPELAEOGRIPHACEA ‣ a fully freshwater order with (1:3/)4 spp. in Spelaeogriphidae, in WC Brazil (Potiicoara), NW Australia (Mangkurtu), and South Africa (Spelaeogriphus) [2]; Brazilian member is the obligatory subterranean species Potiicoara brasiliensis, known only caves from Lagoa Azul in Bonito municipality, Gruta Ricardo Franco near Corumbá municipality, and Gruta do Curupira in the Araras Mountains, Mato Grosso state (Vanin, Zootaxa, 2012).
7. MYSIDA ‣ more than 25 spp. in subterranean environments [2]; 7 cave Mysida occur in Mexico [5], in three families with a single genus each; cave species absents in Brazil and U.S.A.
8. DECAPODA ‣ six infraorders of Decapoda has cave species: Stenopodidae, it is represented by a species of the Macromaxillocarididae family from a anchialine cave in Bahamas; the infraorder Gebiidea recently described another anchialine species of the genus Naushonia (Laomediidae) also for the Bahamas; infraorder Caridea includes a huge of shrimps; Astacidae infraorder with more than 40 stygobitic species of crayfishes of different genera and in environments of temperate and tropical freshwater caves, for Papua New Guinea, Cuba, Mexico, and North America; troglobitic Anomurans occur in Canary Is. (Munidopsis polymorpha), and in SE Brazil (Aegla); Brachyura has troglobitic crabs in Pseudothelphusidae and Trichodactylidae [2].
China has 23 spp., and U.S.A c. 40 of them [2]. Only four in Brazil, all Anomurans in family Aeglidae. Mexico has 27 spp. of troglobitic decapoda, being 17 shrimps: Agostocarididae (1), Alpheidae (3), Barbouridae (1), Hippolytidae (1), Palaemonidae (11), Procarididae (1); 5 crayfishes: Cambaridae (5); and 5 crabs: Pseudothelphusidae (4) and Trichodactylidae (1). U.S.A. has (3:7/)40 spp. [2], 35 crayfishes of Cambaridae, two shrimps of Atyidae, and three shrimps of Palaemonidae. Venezuela includes two troglobitic crabs in Pseudothelphusidae: Chaceus caecus and Eudaniella sp. (Galán & Herrera, BSVE, 2006).
AEGLIDAE
9. MICTACEA ‣ a monotypic order, described for marine caves of Bermuda [2].
10. ISOPODA ‣ more than 300 troglobiotic species in 16 families of terrestrial isopods are known, mostly from the northern part of the globe; 18 in China and (6:18/)104 in U.S.A. [2]. In Brazil, (11:)105 are considered to be troglobionts by our list in at least 21 genera. Mexico has (11:)52 spp. (11 terrestrial and 41 aquatic) of troglobitic Isopoda. Colombia includes a single Armadillidium sp. (Angarita-Sierra, CCE/2018). Venezuela includes six troglobitic Isopoda in six families: Anthuridae (a anchialine species), Calabozoidae, Cirolanidae, Sphaeroniscidae, Oniscidae and Pillosciidae (Galán & Herrera, BSVE, 2006).
45 spp. remains undescribed in Brazil, in Tocantins [TO/1], Rio Grande do Norte [RN/1], Minas Gerais [MG/1][MG/4][MG/6][MG/10], Bahia [BA/1][BA/4], Goias [GO/1], São Paulo [SP/2], Espirito Santo [ES/1] and Mato Grosso do Sul [MS/1].
ONISCOIDEA
ARMADILLIDAE
BRASILEIRINIDAE
CALABOZOIDAE
CIROLANIDAE
DUBIONISCIDAE
PHILOSCIIDAE
PLATHYARTHRIDAE
PUDEONISCIDAE
SCLEROPACTIDAE
STYLONISCIDAE
TRACHELYPODIDAE
11. BOCHUSACEAE ‣ deep sea benthos, but includes two stygobitic species of anchialine caves of the Bahamas. [2].
12. TANAIDACEA ‣ marine benthic habitats around the world, inhabiting anchialine and marine caves in different regions of the world such as Bermuda and several islands of the South Pacific [2].
13. CUMACEA ‣ this group in general has a worldwide distribution with more than 800 recognized species, but subterranean species are collected from Bermudas, Bahamas and Jamaica [2].
5.19 ARTHROPODA/BRANCHIOPODA ‣ troglomorphic forms occur only in Europe, specifically Spain, Romania, France, Slovenia, Bosnia and Herzegovina, all in Diplostraca [2].
5.20 ARTHROPODA/REMIPEDIA ‣ all members of this class are cave-dwelling species in marine anchialine [2], including the two species of Mexico [2, 5].
5.21 ARTHROPODA/CEPHALOCARIDA ‣ none cave dwellling members.
5.22 ARTHROPODA/COLLEMBOLA ‣ c. 500 troglobitic species [4], (7:15/)97 in North America [2], 42 in China [2], also (9:)42 in Mexico [5], and (6:9/)84 spp. in Brazil. All orders of Collembola has troglobites in Brazil except Neelipleona (with 15 troglobitic species, all in Europe and Asia), by Zeppelini (Pre Print, 2021). Besides citation in [4], neither Mexico neither New World no has true troglobitics in order Neelipleona. Arrhopalites mendoncae, Coecobrya phoenix, Pararrhopalites queirozi are cited as troglobitic in [6], none of them is truw cave-adapted (Brito et al., Zootaxa, 2019). Onychiurus acuitlapanensis (Onychiuridae) is a unqiue troglobitic Collembola from Venezuela, known also from Guerrero state in Mexico (Galán & Herrera, BSVE, 2006).
UNDEFINED ORDER ‣ a species identified in N Minas Gerais [MG/5] cannot be detected at the order level.
UNDEFINED SYMPHYPLEONA ‣ a species of this order identified in SE Bahia [BA/1] cannot be detected at the family level; all troglobitic members of this order (c. 100, [4], this source) belongs Arrhopalitidae and Sminthuridae, both present in Brazilina caves. Mexico has 4 troglobitic spp. in both families.
SYMPHYPLEONA/ARRHOPALITIDAE ‣ 14 troglobitic spp. in Brazil and 31 in North America; especially diversified in temperate regions of northern hemisphere, troglobites of this family are present in South America and SE Asia, but absent in Africa; undescribed species in Bahia [BA/2][BA/4], Minas Gerais [MG/3][MG/10] and Rio Grande do Norte state [RN/1].
SYMPHYPLEONA/SMINTHURIDAE ‣ cave species widely in southen Hemisphere, with Adelphoderia in Australia, Troglospinotheca in Argentina, and Pararrhopalites in the tropics, all poorly known taxonomically. Three undecribed species, in Bahia [BA/1] and in Minas Gerais [MG/3] states.
PODUROMORPHA/HYPOGASTRURIDAE ‣ cave species exist in four of the nine families of Poduromorpha order: Gulgastruridae, Hypogastruridae (unique with troglobitics in Brazil), Onychiuridae, and Neanuridae; troglobitic Hypogastruridae of oligotrophic habitats are often characterized by slender or extremely slender claw, but elongation of appendages is moderate; guanobitics have usually reduced eyes and pigment and sometimes elongated claw like several Pseudacherontides. Two undescribed species in Minas Gerais state [MG/2][MG/10]. Mexico has 11 troglobitic species in order, at 4 families.
ENTOMOBRYOMORPHA UNPLACED FAMILY ‣ Entomobryomorpha order is constituted by nine extant families, six of them include cave species; two species of this order identified in SE Bahia [BA/1] and one in São Paulo [SP/3] cannot be detected at the family level. Mexico has 27 spp. in 4 families.
ENTOMOBRYOMORPHA/ISOTOMIDAE ‣ a undescribed species in Brazil, from São Paulo state [SP/2].
ENTOMOBRYOMORPHA/CYPHODERIDAE ‣ are all phyletically blind and unpigmented; most are myrmecophilous or termitophilous, but Cyphoderus and Troglobius have troglobitic-guanobitic species in tropical caves. None exhibits appendage or claw elongation. 10 undescribed troglobitic spp. in Brazil, in Mato Grosso do Sul [MS/1], Minas Gerais [MG/1], Goiás [GO/1] São Paulo [SP/2] and Rio Grande do Norte [RN/1].
ENTOMOBRYOMORPHA/ENTOMOBRYIDAE ‣ subterranean radiations are known for many genera, in various regions of the world. Six troglobitic species undescribeds in Rio Grande do Norte [RN/1] Bahia [BA/3], Mato Grosso do Sul [MS/1] and Minas Gerais [MG/3] states.
ENTOMOBRYOMORPHA/PARONELLIDAE ‣ this family is the most diversified Collembola in many tropical caves but are absent in large areas like southern Sunda Islands. Undescribed troglobitic species in Brazil found in Mato Grosso do Sul [MS/1], Bahia [BA/3], Goiás [GO/1], Rio Grande do Norte state [RN/1], Minas Gerais [MG/3][MG/5][MG/10] and São Paulo [SP/2] states.
5.23 ARTHROPODA/DIPLURA ‣ following Sendra et al. (Zoological Journal of Linnean Society, 2021), 153 described spp. are obligate-subterranean, 110 in Europe to adjacent islands, Magreb (3), Lebanon (1), Turkmenistan (1), South Africa (1), India (1), Myanmar (1), China (5), Japan (1), Indonesia (1), Papua New Guinea (2), Australia (1), Canada (one Haplocampa), U.S.A. (13, Eumesocampa, Haplocampa, Condeicampa and Mixojapyx one each, and nine Litocampa), Mexico (two Litocampa, two Juxtlacampa, one Oncinocampa and three Tachycampa), Guatemala (one Juxtlacampa) and one in Brazil, Oncinocampa trajanoae Condé, from Alto do Ribeira karst, São Paulo state (this species is corroborated by [6]).
Nine undescribed species was collected in Brazil, in Rio Grande do Norte [RN/1], Minas Gerais [MG/1][MG/2] and Bahia [BA/2][BA/3][BA/5].
UNDETERMINED FAMILY
CAMPODEIDAE
JAPYGIDAE
PROJAPYGIDAE
5.24 ARTHROPODA/PROTURA ‣ unknown obligate subterraneans.
5.25 ARTHROPODA/INSECTA ‣ members of 11 orders of insects occur as troglobitic, by [4]. 290 troglobitic insects occur in North America: Coleoptera (269), Dermaptera (1), Diptera (1), Hemiptera (9), Orthoptera (3), Zygentoma (7); 202 in China [2]. True troglobitic insecta in Mexico are 65 spp. in 4 orders: Zygentoma (13), Orthoptera (4), Hemiptera (3, all Homoptera) and Coleoptera (43). Brazil has 83 spp. in eight orders: Zygentoma (6), Orthoptera (4), Hemiptera (7), Coleoptera (36), Blattodea (5), Diptera (1), Dermaptera (1) and Hymenoptera (2). Colombia possibly includes one troglobitic Ephemeroptera (Angarita-Sierra, CCE/2018, also Angarita-Sierra, CCE/2018; if confirmed, will be the first troglobitc in this order worldwide).
A. GRYLLOBLATTODEA ‣ despite numerous citations of members of this order in caves (7 troglobitic species by [4]), nothing about troglomorphic forms is available in the literature.
B. ZYGENTOMA ‣ c. 45 troglobitics are known at present, most belong to Nicoletiidae, except two Protrinemuridae from Greece and Thailand. Nicoletiidae is subdivided into four subfamilies: Atelurinae (a single species from Philippines), Coletiinae (12 troglobites in the Mediterranean basin, South Africa, Brazil and Thailand), Subcoletiinae (3 troglobites in Southeast Asia, one in New Guinea and one in Australia), and Cubacubaninae (25 troglobites distributed from Central America to southern USA) [4]. 7 spp. in U.S.A. (6 Texoreddellia and one Speleonycta) and 13 in Mexico (in three genera). 7 spp. in Brazil, one described and six undescribeds in Minas Gerais [MG/1], Bahia [BA/2][BA/3][BA/4] and Espírito Santo [ES/1], inc. two in Lepismatidae; if published, these will be the first troglobitic species in this family.
UNDETERMINED FAMILY
LEPISMATIDAE
NICOLETIIDAE
C. ORTHOPTERA ‣ all cave species of Orthoptera are members of Ensifera, which are omnivorous or scavengers, and none of Caelifera, which are phytophagous; most belong to the large families Gryllidae, Phalangopsidae, Trigonidiidae, and Rhaphidophoridae. A noticeable exception is the two troglobitics Rhaphidiophoridae from Venezuela, which are the only aquatic or semi-aquatic cave Orthoptera (Galán & Herrera, BSVE, 2006). c. 250 troglobitic species worldwide, [4], only 4 in Brazil, all in Phalangospsidae (inc. one undescribeds in Bahia [BA/4]). Some troglophylic Phalangopsis (Phalangopsidae, Junta et al., Zootaxa, 2020) occur in Amazon region. All six troglobitic Orthoptera from Mexico are also Phalangospsidae; all three U.S.A. members are Gryllidae from Hawaii [2].
PHALANGOSPSIDAE
D. COLEOPTERA ‣ the most speciose order of life, with c. 390,000 spp.; c. 2,500 spp. are troglobitic; three families, Carabidae, Staphylinidae (including Pselaphinae and Scydmaeninae), and Leiodidae count for 98% of the diversity of troglobitic and troglophilic beetles, with 1180, 110, and 599 spp., respectively [4]. Mexico has 44 spp. in Carabidae (30), Dytiscidae (1), Histeridae (4), Leiodidae (8) and Ptinidae (1); Colombia has 5 troglobitic Coleoptera (two in Staphylinidae; Carabidae, Nitidulidae, Passalidae one each; Angarita-Sierra, CCE/2018); U.S.A. has (9:31/)269 cave-obligate species, 147 in Pseudanophthalmus. Venezuela includes three troglobitics in Carabidae, Dysticidae and Catopidae (Galán & Herrera, BSVE, 2006).
47 in Brazil (23 undescribeds in Bahia [BA/1][BA/2][BA/4][BA/5], Mato Grosso do Sul [MS/1], Goiás [GO/1], Minas Gerais [MG/1][MG/2][MG/6][MG/10], Rio Grande do Norte [RN/1], São Paulo [SP/2], Tocantins [TO/1]), in three families. Truly troglobitic species at Scarabeoidea are yet to be found in Brazil (C.M.A. Correa et al., Revista Brasileira de Entomologia, 2022).
CARABIDAE
DYTISCIDAE
STAPHYLINIDAE
E. BLATTODEA ‣ about 40 troglobitic species, all saprophagous and from Old World, except three fully described in New World, Ischnoptera peckorum from Galapagos Is., Ecuador (Ectobiidae), Paranocticolla venezuelana from Venezuela (Galán & Herrera, BSVE, 2006), and Litoblatta camargoi (Blattellidae), from Bahia state (Guterrez, Solenodon, 2005); additional five undescribed troglobitic species are found in Brazil, in Bahia [BA/1][BA/2][BA/3] and Minas Gerias [MG/2][MG/6], and one in Colombia (Angarita-Sierra, CCE/2018).
F. PSOCOPTERA ‣ besides data in [4] for several possible troglobitic in New World, the only clearly troglobitic representative of this order in New World is the prionoglaridid Speleopsocus chimanta known from a humid cave in SE Venezuela (C. Lienhard & R. L. Ferreira, Revue suisse de Zoologie, 2015).
G. DERMAPTERA ‣ there are only two troglobites with well-marked troglomorphic traits in this order, Anataelia troglobia (Canary Is.) and Anisolabis howarthi (Hawaii) [4]; in Brazil the cave species Mesodiplatys falcifer from Carinhanha (Diplatyidae, Bahia) is possibly troglobitic (Kamimura, ZooKeys, 2018) and accepte here; this species is not accepted as troglobitic by [6]. One possibly troglobitic species was collected in Colombia (Angarita-Sierra, CCE/2018).
H. HEMIPTERA ‣ 85 spp. are caverniculous worldwide [4]. Brazil has 12 troglobitic spp. in 6 families in all three suborders with cave members. All three Mexican troglobitics of this order belongs Fulguromorpha, and all five troglobitics in U.S.A. are endemic to Hawaii, in Cixiidae (Fulguromorpha) and Reduviidae (Heteroptera). One possibly troglobitic species in Cydnidae was collected in Colombia (Angarita-Sierra, CCE/2018).
Auchenorrhyncha/Fulguromorpha ‣ c. 60 spp. in four families are troglobitics: Cixiidae, Meenoplidae, Kinnaridae, and Delphacidae. Brazil has six spp. in three of these families, one undescribed in Bahia [BA/5].
CIXIIDAE
DELPHACIDAE
KINNARIIDAE
Heteroptera ‣ c. 50 spp. a small proportion of them are found in subterranean habitats, as guanophiles (Leptopodidae, Lygaeidae, and Reduviidae), bat ectoparasites (Cimicidae), or troglophiles and troglobites of parietal associations (Reduviidae, Emesinae). Brazil has 5 spp. in two families, two undescribed in Mato Grosso do Sul [MS/1] and Bahia [BA/5] states.
DIPSOCORIDAE
HYDROMETRIDAE
Stenorhyncha ‣ none informations, but one undescribed was collected in Brazil, in Ortheziidae.
ORTHEZIIDAE
I. DIPTERA ‣ the discussion about troglobitic diptera is confusing because all troglobitic members of this order are not or weakly modified for cave life; several non-hematophagous Diptera are troglobitics, c. 20 spp., all from Old World except Spelobia tenebrarum, the only troglobitic Diptera of North America; the only hematophagous species firmly established as a troglobite, Deanemyia maruaga from Brazil (known only from Maruaga cave, in Presidente Figueiredo, Amazonas state), is parthenogenetic, nonvector, and can complete its full life cycle in cave [4]; see Veracilda Ribeiro Alves et al. (Mem. Inst. Oswaldo Cruz, 2011); this species is corroborated by [6]. Two possibly troglobitic species was collected in Colombia (Drosophilidae and Mycetoohilidae one each, Angarita-Sierra, CCE/2018).
J. LEPIDOPTERA ‣ only two described troglobitic species in over order: the reduced-eyes Tineidae, Tinea microphthalma in Philippines and the Erebidae Schrankia howarthi of Hawaii, which presents several regressive traits: flightlessness, reduced pigmentation, and reduced eye size [4]. One possibly troglobitic species was collected in Colombia (Angarita-Sierra, CCE/2018).
K. HYMENOPTERA ‣ three described members of this order, all ants, are fully troglobitics: Leptogenys khammouanensis, collected in oligotrophic habitats of a Laotian cave, is possibly the only troglobitic ant [4]; Aphaenogaster gamagumayaa, from Okinawa, Japan (Naka & Maruyama, Zootaxa, 2018); and Yavnella laventa Griebenow, Moradmand & Isaia, known from Fars province, SW Iran (Z. H. Griebenow et al., Invertebrate Systematics, 2022). Two undescribed species was collected in Bahia [BA/1] and Tocantins [TO/1] states, Brazil.
5.26 PHYLUM CRANIATA/PISCES ‣ 280 spp. among 11 orders, by Subterranean Fishes of the World (SFW): Characiformes only in Mexico and Brazil; Cyprinodontiformes only in Mexico; Gymnothiformes only in Brazil; Percopsiformes and Perciformes only in U.S.A; Siluriformes in Asia, Africa and New World; Ophidiformes in Caribbean, Mexico, Galapagos and Indonesia; Gobiiformes in Japan, Madagascar, Australia, Guam, Mexico, Indonesia, Philippines and Papua New Guinea; Synbrachiformes in India, Australia, Cameroon, Mexico and Venezuela; Anabanthiformes only in India; and Cypriniformes in Germany, Africa and Asia from Iraq and Turkmenisthan to Malaysia and China.
By SWF, Brazil appears with 37 spp., in secord position, below China (88 spp., 86 only in Cypriniformes, mainly in Yunnan region), with 19 spp. more than Mexico and 15 more than the U.S.A. The four countries together have 164 spp. (U.S.A. and Mexico shared one spp.), more than half of the world amount of species.
Neolissochilus pnar (Cyprinidae) endemic to NE India is the largest troglobtic fish worldwide, possibly the largest of all troglobites (Novotaxa).
BY COUNTRY
Brazilian diversity is 36 spp. of 12 genera in 7 families of three orders, inc. the unique cave fish at order Gymnotiformes (Eigenmannia vicentespelea); also Stygichthys (Characidae), Aspidoras (Callichtyidae), Ituglanis, Glaphyropoma, Copionodon, Trychomycterus (Trichomycteridae), Ancistrus, Isbrueckerichthys (Loricariidae), Phreatobius, Pimelodella, Rhamdia, and Rhamnidiopsis (Heptapteridae). All species mentioned in SFW are on [1] list except Isbrueckerichthys alipionis and Phreatobius sanguijuela. Venezuela includes 7 troglobitic fishes in Trichomycterus (2); Ancistrus (1); Synbranchus (1) and Rhamdia (1), by Galán & Herrera, BSVE, 2006.
Mexico has (8:8/)13 spp. in 5 orders, inc. the unique cave fish at order Cyprinodontidae (Poecilia mexicana); also Rhamdia (Heptapteridae), Astynax (Characidae), Prietella (Ictaluridae), Lucifuga (Bithytidae), Typhlias (Dinematichthyidae), Caecieleotris (Eleotridae) and Ophisternon (Synbranchidae).
In the U.S.A. there are (3:7/)21 spp. in 3 orders, in genera Prietella, Satan, Trogloglanis (Ictaluridae), Amblyopsis, Speoplatyrhinus, Troglichthys (Ambryopsidae) and Cottus (Cottidae).
REGIONAL
All genera with troglobite fish in South America have them in Brazil except Astroblepus (Ecuador and Peru one troglobitic each, Astroblepidae), Chaetostoma (1 troglobitic, in Ecuador, Loricariidae), Silvinichthys (1 troglobitic, in Argentina, Trichomycteridae), Ogilbia (1 troglobitc, Galapagos, Dinematichthyidae) and Synbranchus (1 troglobitic, Venezuela), all of them with troglobiotics only on the continent.
In New World, exclusively underground endemic genera are Trogloglanis and Satan, both monotypic of Ictaluridae endemic to the U.S.A.; Amblyopsis, Triglichthys and Speoplatyrhynus, all of Amblyopsidade, which is, as a whole, an endemic family of the U.S.A. No other country has entirely underground endemic genera of fish. In hemisphere only Prietella phreatophila (Mexico/U.S.A.) and Phreatobius sanguijuela (Brazil/Bolivia) are not national endemisms. Altogether there are 95 spp. in 12 countries, distributed in 30 genera of 16 families in 9 orders.
OUR LIST
In our list, we recognied all species cited in [1] plus three undescribed in Mato Grosso do Sul [MS/1] and Bahia [BA/4], and Phreatobius sanguijuela; all troglobite species of fishes cited here are corroborated by [6]. The range of P. cisternarum has been expanded with records from Muriel-Cunha & Pinna (Papéis Avulsos de Zoologia, 2005).
CHARACIFORMES ‣ troglobitic in this order includes only three Characidae: two Astynax from Mexico and the Brazilian Stygichthys.
GYMNOTIFORMES ‣ Brazilian cave Eigenmannia (Sternopygidae) is the unique stygobitic in this order worldwide.
SILURIFORMES ‣ order with 82 cave species in 11 families, 5 of them with cave members in Brazil.
CALLYCHTHYIDAE
HEPTAPTERIDAE
LORICARIIDAE
TRYCHOMYCTERIDAE
5.27 PHYLUM CRANIATA/AMPHIBIA ‣ by [2], 12 (13, mentioned in work, is a mistake for source) amphibians worldwide are obligatory subterranean species: 11 Pletodontidae from U.S.A., being 8 Eurycea (4 in Texas, 3 in Ozark karst area in Arkansas, Kansas, Missouri, Oklahoma; one in Florida and Georgia), and three Gyrinophilus (Alabama, Georgia, Tennessee and West Virginia); and Proteus anguinus (Proteidae), the only stygobiotic salamander in Europe and unique member of their family in continent, reaches a length of more than 25 cm, making the second largest stygobiont (troglobitic) known anywhere. It occurs throughout the Dinaric Mountains in northeast Italy, Slovenia, Croatia, and Bosnia-Herzegovina. [5] does not recognize any cave salamander in Mexico.
LAST UPDATES
26∙04∙2024 ‣ exclusion of Endecous apterus and E. peruassuensis (Orthoptera), rejected in Carvalho, P.H.M. et al (Zootaxa, 2023).
25∙04∙2024 ‣ addition of 11 new described species: Pinelema elinae (BA), Loxosceles boqueirao (BA), L. bodoquena (MS), Spaeleoleptes gimli (BA), Cayenniola albaserrata (BA), Trichorhina baiana (BA), Ctenorillo iuiuensis (BA), Circoniscus mendesi (PA), Circoniscus xikrin (PA), Kadiweuoniscus rebellis (MS) and Endecous vitreus.
22∙01∙2024 ‣
inclusion of 10 new undescribed species in Aguas Claras System, Carinhanha (BR), in Araneae, Coleoptera, Diplura, Hemiptera and Dilplopoda (SEE).
22∙01∙2024 ‣ exclusion of Pseudonannolene canastra (Diplopoda), synonimized under P. ambuatinga, with ratification of accounting on several topics and readaptation of the P. ambuatinga map (SEE).
20∙10∙2023 ‣ inclusion of Caraiboscia jabutiensis (Isopoa, MT, SEE), Gabunillo enfurnado, Venezillo moreirai, V. limai (Isopoda, BA, SEE) and Spelaeometra hypogea (Hemiptera, BA, SEE).
16∙10∙2023 ‣ of records of undescribed troglobite species in three regions: Lapa Nova, Vazante, MG (6 spp., SEE), Lagoa Santa, MG (4 spp., SEE) and Presidente Olegário, MG (3, SEE).
10∙09∙2023 ‣ updates in undescribed species in Bahia state, with a inclusion f 30 news undescribeds (Gallão et al., Diversity, 2023); inclusion of Plutogeophilus jurupariquibaba, former undescribed (Chilopoda, [SP/2], SEE); inclusion of Benthana alba, Benthanoides amazonicus, B. tarzan (Cardoso & Ferreira, Zootaxa, 2023), Ferricixius michaeli and F. goliathi (Ferreira et al., Zootaxa, 2023).
25∙07∙2023 ‣ updates in troglobitic Hirundina and Polychaeta at Annelida.
20∙07∙2023 ‣ updates in Amblypygi, with a remarkable new record in Colombia.
30∙06∙2023 ‣ updates about troglobitic sponges, and troglobits in Mexico, and inclusion of Phretobius sanguijuela, with general updates in Heptapteridae.
19∙06∙2023 ‣ inclusion of Endecous infernalis, former undescribed [BA/1](SEE).
09∙06∙2023 ‣ add new species: Pseudochthonius koinopoliteia former undescribed [BA/], P. diamachi, P. pali (Prado & Ferreira, Zootaxa, 2023).
03∙01∙2023 ‣ updates in Collembola, inclusion of Iandumoema cuca, I. gollum, I. stygi (Opiliona), Dobrodesmus mirabilis (Dobrodesmidae), Ardistomis ferreirai (Carabidae), Metopioxys carajas, Oxarthrius aurora, O. inexpectatus (Staphylinidae), Leptus sidorchukea (Erythraeidae), Tonton itabirito, T. matodentro (Microstigmatidae), Pararrhopalites sideroicus (Sminthuridae), Onciurosoma troglobium (Paradoxosomatidae), Loxocelis troglobia (Sicariidae), Brasilomma enigmatica, Indiani gaspar, Paracymbiomma bocaina, P. pseudocaesus (Prodidomidae), Ochyrocera dorinha, O. rosinha, O. ungoliant, Speocera babau, S. pinima, S. piquira (Ochyroceratidae), Charinus cearensis, C. diamantinus, C. euclidesi, C. puri, C. renneri (Charinidae), Aegla charon (Aeglidae), Lavajatus moroi (Subulinidae), and exclusion of Troglorhopalurus lacrau (Scorpiona), Isoctenus corymbus (Ctenidae), Lygromma ybyguara (Prodidomidae) and Spiripockia umbraticola (Pomatiopsidae), by indications in [6], and inclusion of Eukoenenia mocororo.
08∙08∙2022 ‣ conclusion of a huge revision of all data from this page, inc. several notes for Mexico and U.S.A.
03∙07∙2022 ‣ a huge revision, including c. 280 new species in list, mainly undescribed.
12∙06∙2022 ‣ inclusion of Perigona spelunca (Carabidae).
26∙11∙2021 ‣ inclusion in the list of troglobionts in Brazil of 6 spp. de Matta (Araneae) and updated the text to include this addition.