Milyeringa veritas

Whitley 1945

Milyeringa veritas
Milyeringa veritas. Photo used with permission of David Morgan


Photo of Milyeringa veritas by David Morgan and used with permission


Milyeringa brooksi  Chakrabarty 2010

Synonymised with Milyeringa veritas by Larson et al. (2013).




Holotype: WAM P.2913-001, 40.5mm SL. This species is the type species by original designation and monotypy of the cave restricted genus Milyeringa, and the genus is the type genus of the family Milyeringidae.


Type locality: Milyering Well (=C-24), near Yardie Creek Station, North West Cape, Western Australia (22o01’S, 113o56’E). Details of all other known locations are given in Humphreys (1999) and summarised below. One further site is given in Page et al. (2018). At ten locations within Cape Range this species has been seen at sites from which Ophisternon candidum has also been recorded (Humphreys and Adams 1991, Adams and Humphreys 1993, Humphreys 1993, Knott 1993). The range of Ophisternon candidum has recently been extended to the east of the Pilbara region, but to date M. veritas has not been found in this area (Moore et al. 2018).

Area Site No. Site name Species Latitude, longitude Clade References
CRE C-27 Kubura Well Mv, Oc


veritas Humphreys (1999)
CRE C-495 New Mowbowra Cave Mv, Oc
-21.9833 114.1167
veritas Humphreys (1999)
CRE C-361 Mowbowra Well Mv, Oc
-21.9933 114.1206
veritas Humphreys (1999)
CRE C-23 Dozer Cave Mv, Oc   veritas? Humphreys (1999)
CRE C-282 Ned's well Mv   veritas? Humphreys (1999)
CRE C-452 Cameron's Cave Mv   veritas? Humphreys (1999)
CRE WC44 None Mv   veritas? Humphreys (1999)
CRW C-414 Wobiri Rock Hole Mv, Oc
-21.8333 114.0667
veritas Humphreys (1999)
CRW C-273 Five Mile Well Mv
-21.8500 114.0650
veritas Humphreys (1999)
CRW None Ampolex D Mv
-21.8632 114.0286
vertitas Page et al. (2018)
CRW C-25 Kudumurra Mv, Oc
-21.8880 114.0090
veritas brooksi Humphreys (1999)
CRW C-24 Milyering Well Mv Oc
-22.0200 113.9295
veritas Humphreys (1999)
CRW C-149 Tulki Well Mv
-22.0920 113.8970
brooksi Humphreys (1999)
CRW C-274 Pilgramunna Well Mv, Oc
-22.1920 113.8660
brooksi Humphreys (1999)
CRW C-28 Bundera Mv   brooksi Humphreys (1999)
CRW C-362 Javis Well Mv
-22.5967 113.6906
brooksi Humphreys (1999)
CRW C-215 Unnamed Mv, Oc 22o2'S 113o56' E brooksi?

Humphreys (1999)

Moore et al (2018)

CRW C-332 Tantabiddi Rockholes Mv   brooksi? Humphreys (1999)

The distribution of Milyeringa veritas. Mv Milyeringa veritas, Oc Ophisternon candidum, CRE Cape Eange East, CRW Cape Range West


Various stygobitic Crustacea are found in Cape Range, eastern Pilbara and Barrow Island and their biogeography can help to inform us about the distribution and ages of the fish taxa there. The following sources are useful in this regard:

Humphreys (1993) Stygofauna and a Tethyan connection

Yager and Humphreys (1996) Lasionectes exleyi (Crustacea: Remipedia)

Page et al. (2008) Stygiocaris (Decapoda: Atyidae)

Page et al. (2018) Halosbaena (Peracarida: Thermosbaenacea)

Matthews et al. (2019) Parabathnellidae diversity in Pilbara


Found in both freshwater and anchialine systems from close to the coast up to 4.3 km inland. Seen at karst windows and a depths of up to 33m. Collected from boreholes very remote from any opening to the surface (details from Humphreys 2001). The food of this species (and of Ophisternon candidum) was examined by Humphreys and Feinberg (1995). They found that 79% of the intake was terrestrial fauna in the form of Isopoda, Trichoptera, Hymenoptera and cockroaches. Only 10% maximum intake was from consumption of other cave animals. The Cape Range karst is the 4th most biodiverse subterranean habitat in Australia with 83 obligate subterranean species (41 terrestrial and 42 aquatic). The Barrow Island Karst is the 5th most diverse with 74 obligate species (18 terrestrial and 56 aquatic) (Ebergard and Haworth 2021).


Whitley (1945) erected the family Milyeringidae to contain this species because it differed from the Eleotridae by having no eyes and reduced ventral and dorsal fins. Subsequent authors (e.g. Greenwood Rosen, Weizman and Myers 1966, Nelson 1994) have not recognised this family and have placed the species in the Eleotridae. Hoese and Gill (1993) made a thorough study of eleotridid fishes within the family Gobiidae and placed the genus Milyeringa in the subfamily Butinae together with the genus Typhleotris.

Larson and Foster, in Larson, Foster Humphreys and Stevens 2013, descibed a second species in the genus, Milyeringa justitia, from Barrow Island, offshore from the Western Cape.

Larson et al. (2013) synonymised M brooksi with M. veritas but there is a clear genetic and geographic difference between the two (their Figure 3) with the brooksi clade, concentrated at sites in south west cape Range, sister to the veritas clade, concentrated at sites in north west and east Cape Range, with an overlap only at C-25, Kudumurra Well. A comprehensive molecular analysis by Page et al. (2018), with data from M. veritas, M. brooksi, M. justitia, Typhleotris madagascariensis, T. pauliani and T. mararybe, supported the erection of a new species for jusitia and the synonymisation of brooksi with veritas. However, their data still show a clear sister relationship between the latter two taxa.

Their data also strongly support the placement of both Milyeringa and Typhleotris in the Family Milyeringidae as suggested by Chakrabarty et al. (2012) and Sparks and Chakrabarty (2012).

Kuang et al. (2018) used molecular markers to study the internal structure of the Suborder Gobioidei which contains within it eleven Families. They demonstrated that Milyeringidae is sister Family to a group of four  Families: Oxudercidae+Gobiidae+Butidae+ Eleotridae and is only distantly related to four others: Odontobutidae, Rhyacichthyidae, Apogonidae and Kurtidae (samples from Thalasseleotridae and Trichonotidae were not availabe for analysis).

Conservation Status

MuG [EN B1ab(ii,iii)+2ab(ii,iii): 3.1: 2019]

Moore, G. 2019. Milyeringa veritas. The IUCN Red List of Threatened Species 2019: e.T13555A123378349.

Wager, R. 1996. Milyeringa veritas. The IUCN Red List of Threatened Species 1996: Downloaded on 05 July 2017

R (IUCN 1994), DD (IUCN 1996, 2000). Rare (Michaelis 1985). Humphreys and Adams (1991) document an incursion of saline water into the freshwater aquifer which, if real and continuous, will represent a dangerous threat to this and other stygobitic species in the coastal fringe of the Cape Range. This fringe is outside of the Cape Range National Park and therefore does not get the protection offered to sites within it.

From the pont of view of conservation the principle of precaution should hold and if this is done in this case the clade brooksi should be considered, at minimum, as an evolutionary significan unit (ESU) and not sunk entirely into the veritas clade ss.

The conservation of karst sites, and their fauna, in western Autralia is considered in a very detailed report by the Environmental Protection Authority (2012).


Museum Holdings

26 specimens at the Western Australian Museum, Perth (Whittaker and Kritski 1973), AMS I.21336-001 4 specimens 31-43mm SL (Hoese and Gill 1993), AMNH 45497, AMNH 48568 (Humphreys and Feinberg 1995).

Internet Resources

Ningaloo Coast world heritage nomination with details of the subterranean fauna

Key References

Whitley, G. P. Journal Article 1945 New sharks and fishes from Western Australia. Part 2
Holthuis, L. B. Journal Article 1960 Two new species of Atyid shrimps from subterranean waters of N.W. Australia (Decapoda, Natantia)
Mees, G.F. Journal Article 1962 The subterranean freshwater fauna of Yardie Creek Station, North West Cape, Western Australia
Cawthorn, P. Journal Article 1963 Discovery of subterranean freshwater fauna on the eastern side of North West Cape
Richards, A. M. Journal Article 1963 The subterranean freshwater fauna of North West Cape, Western Australia
Whittaker, F. H. and Kritsky, D. C. Journal Article 1973 An examination of the Australian blind cavefish Milyeringa veritas Whitley, 1945, for helminth parasites
Merrick, J. R. and Schmida, G. E. Book 1984 Australian freshwater fishes: Biology and management
Michaelis, F. Journal Article 1985 Threatened fish. A report on the threatened fish of inland waters of Austalia
Young, M. Journal Article 1986 Bringing the blind gudgeon into captivity
Allen, G.R. Book 1989 Freshwater fishes of Australia
Pollard, D. A., Ingram, B. A., Harris, J. H. and Reynolds, L. F. Journal Article 1990 Threatened fishes in Australia - an overview
Humphreys, W. F. and Adams, M. Journal Article 1991 The subterranean aquatic fauna of the North West Cape peninsula, Western Australia
Allen, G.R. Book 1992 Inland Fishes of Western Australia
Hoese, D.F. and Gill, A.C. Journal Article 1993 Phylogenetic relationships of eleotridid fishes (Perciformes, Gobioidei)
Adams, M. and Humphreys, W.F. Journal Article 1993 Patterns of genetic diversity within selected subterranean fauna of the Cape Range penninsula, Western Australia: Systematic and biogeographic implications
Knott, B. Journal Article 1993 Stygofauna of the Cape Range penninsula, Western Australia: Tethyan relicts
Humphreys, W. F. Book 1993 The biogeography of Cape Range, Western Australia
Humphreys, W. F. Journal Article 1993 The significance of the subterranean fauna in biogeographical reconstruction: Examples from Cape Range Peninsula, Western Australia
Humphreys, W.F. Journal Article 1994 The subterranean fauna of the Cape Range coastal plane, Northwestern Australia
Humphreys, W. F. and Feinberg, M. N. Journal Article 1995 Food of the blind cave fishes of Northwestern Australia
Yager, J. and Humphreys, W. F. Journal Article 1996 Lasionectes exleyi sp. nov. the first Remipede crustacean recorded from Australia and the Indian Ocean, with a key to the world species
Eberhard, S. and Hamilton-Smith, E. Journal Article 1996 Conservation of cave fauna in Australia
Hamilton-Smith, E., Kiernan, K. and Spate, A. Journal Article 1996 Karst management considerations for the Cape Range karst province, Western Australia
Humphreys, W.F. Book Section 1997 Protection of arid karstlands on the Cape Range Penninsula, Western Australia
Humphreys, W.F., Poole, A., Eberhard, S.M. and Warren, D. Journal Article 1999 Effects of research diving on the physico-chemical profile of Bundera Sinkhole, an anchialine remiped habitat at Cape Range, Western Australia
Humphreys, W. F. Journal Article 1999 Physico-chemical profile and energy fixation in Bundera Sinkhole, an anchialine remiped habitat in north-western Australia
Humphreys, W. F. Journal Article 1999 The distribution of Australian cave fishes
Humphreys, W.F. Book Section 2000 Background and glossary
Humphreys, W.F. Book Section 2000 The hypogean fauna of the Cape Range Peninsula and Barrow Island, northwestern Australia
Romero, A. and Vanselow, P.B.S. Journal Article 2000 Threatened fishes of the world: Milyeringa veritas Whitley, 1945 (Eleotridae)
Moore, B.P., Humphreys, W.F., Decu, V. and Juberthie, C. Book Section 2001 Australie
Jaume, D., Boxshall, G.A. and Humphreys, W.F. Journal Article 2001 New stygobiont copepods (Calanoida; Misophrioida) from Bundera sinkhole, an anchialine cenote on north-western Australia
Humphreys, W.R. Journal Article 2001 Groundwater calcrete aquifers in the Australian arid zone: the context to an unfolding plethora of stygal biodiversity
Wilson, R.S. and Humphreys, W.F. Journal Article 2001 Prionospio thalajani sp. nov. (Polychaeta: Spionidae) from an anchialine cave, Cape range, north-western Australia
Humphreys, W.F. Journal Article 2001 Milyeringa veritas (Eleotridae), a remarkably versatile cave fish from the arid tropics of northwestern Australia
Humphreys, WF Journal Article 2001 The subterranean fauna of Barrow Island, northwestern Australia, and its environment
Unmack, P.J. Journal Article 2001 Biogeography of Australian freshwater fishes
Boulton, A.J., Humphreys, W,F. and Eberhard, S.M. Journal Article 2003 Imperilled subsurface waters in Ausralia: biodiversity, threatening processes and conservation
Hancock, P.J., Boulton, A.J. and Humphreys, W.F. Journal Article 2005 Aquifers and hyporheic zones: Towards an ecological understanding of groundwater.
Thacker, C.E. and Hardman, M. Journal Article 2005 Molecular phylogeny of basal gobiod fishes: Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei)
Eberhard, S.M., Halse, S.A. and Humphreys, W.F. Journal Article 2005 Stygofauna in the Pilbara region, north-west Western Australia: a review
Humphreys, W.F., Shiao, J.-C., Iizuka, Y. and Tzeng, W.-N. Journal Article 2006 Can otolith microchemistry reveal whether the blind cave gudgeon, Milyeringa veritas (Eleotridae), is diadromous within a subterranean estuary?
Humphreys, W.F. Journal Article 2006 Aquifers: the ultimate groundwater-dependent ecosystems
Seymour, J.R., Humphreys, W.F. and Mitchell, J.G. Journal Article 2007 Stratification of the microbial community inhabiting an anchialine sinkhole.
Finston, T.L., Johnson, M.S., Humphreys, W.F., Eberhard, S.M. and Halse, S.A. Journal Article 2007 Cryptic speciation in two widespread subterranean amphipod genera reflects historical drainage patterns in an ancient landscape
Humphreys, W.F. Journal Article 2008 Rising from down under: developments in subterranean biodiversity in Australia from a groundwater fauna perspective
Page, T.J., Humphreys, W.F. and Hughes, J.M. Journal Article 2008 Shrimps downunder: Evolutionary relationships of subterranean crustaceans from Western Australia (Decapods: Atyidae: Stygiocaris)
Thacker, C.E. Journal Article 2009 Phylogeny of Gobioidei and placement within Acanthomorpha, with a new classification abd investigation of diversification and character evolution
Humphreys, W.F., Watts, C.H.S., Cooper, S.J.B. and Leijs, R. Journal Article 2009 Groundwater estuaries of salt lakes: buried pools of endemic biodiversity on the western plateau, Australia
Collins, L.B. and Stevens, A. Report 2010 Assessment of coastal groundwater and linkages with Ningaloo Reef – final report February 2010
Department of the Environment, Water, Heritage and the Arts Report 2010 Ningaloo Coast World Heritage nomination
Chakrabarty, P. Journal Article 2010 Status and phylogeny of Milyerigidae (Teleostei:Gobiiformes), with the description of a new blind cave-fish from Australia, Milyeringa brooksi, n. sp.
Humphreys, W., Tetu, S., Elbourne, L., Gillings, M., Seymour, J., Mitchell, J. and Paulsen, I. Journal Article 2012 Geochemical and microbial diversity of Bundera Sinkhole, an anchialine system in the eastern Indian Ocean
Abrams, K. M., Guzik, M.T., Cooper, S.J.B., Humphreys, W.F., King, R.A., Cho J.L. and Austin, A.D. Journal Article 2012 What lies beneath: molecular phylogenetics and ancestral state reconstruction of the ancient subterranean Australian Parabathynellidae (Syncarida, Crustacea)
Chakrabarty, P., Davis, M.P. and Sparks, J.S. Journal Article 2012 The first record of a trans-oceanic sister-group relationship between obligate vertebrate troglobites
Sparks, J.S. and Chakrabarty, P. Journal Article 2012 Revision of the endemic Malagasy cavefish genus Typhleotris (Teleostei: Gobiiformes: Milyeringidae) with discussion of its phylogenetic placement and description of a new species
Environmental Protection Authority Report 2012 A review of subterranean fauna assessment in Western Australia. Discussion Paper. February 2012
Humphreys, W.F Book Section 2012 Diversity patterns in Australia
Unmack, P.J. Journal Article 2013 Biogeography
Humphreys, G., Alexander, J., Harvey, M.S. and Humphreys, W.F. Journal Article 2013 The subterranean fauna of Barrow Island, northwestern Australia: 10 years on
Tornabene, L., Chen, Y.G. and Pezold, F. Journal Article 2013 Gobies are deeply divided: phylogenetic evidence from nuclear DNA (Teleostei: Gobioidei: Gobiidae)
Larson, H.K., Foster, R., Humphreys, W.F. and Stevens, M. Journal Article 2013 A new species of blind cave gudgeon Milyeringa (Pisces: Gobioidei, Eleotridaee) from Barrow Island, Western Australia, with a redescription of M. veritas Whitley
Halse, S., Scanlon, M., Cocking, J., Barron, H., Richardson, J. and Eberhard, S. Journal Article 2014 Pilbara stygofauna: deep groundwater of an arid landscape contains globally significant radiation of biodiversity
Morgan, D.L., Unmack, P.J., Beatty, S.J., Ebner, B.C., Allen, M.G., Keleher, J., Donaldson, J.A. and Murphy, J. Journal Article 2014 An overview of the 'freshwater fishes' of Western Australia
Bishop, R.E., Humphreys, W.F., Cukrov, N., Zic, V., Boxshall, G.A., Cukrov, M., Iliffe, T.M., Krsinic, F., Moore, W.S., Pohlman, J.W., and Sket, B. Journal Article 2015 ‘Anchialine’ redefined as a subterranean estuary in a crevicular or cavernous geological setting
Rasoloariniaina, J.R., Ganzhorn, J.U., Riemann, J.C. and Raminosoa, N. Journal Article 2016 Water quality and biotic interaction of two cavefish species: Typhleotris madagascariensis Petit, 1933 and Typhleotris maraybe Sparks and Chakrabarty, 2012, in the Mahafaly Plateau groundwater syste, Madagascar
Humphreys, W. F. Book Section 2017 Australasian subterranean biodiversity
Moore, G.I. Journal Article 2018 New morphological data and live photographs of the rare subterranean blind cave eel Ophisternon candidum (Synbranchidae) from north‑western Australia
Moore, G.I., Humphreys, W.F. and Foster, R. Journal Article 2018 New populations of the rare subterranean Blind Cave Eel Ophisternon candidum (Synbranchidae) reveal recent historical connections throughout northwestern Australia
Page, T.J., Hughes, J.M., Real, K.M., Stevens, M.I., King, R.A. and Humphreys, W.F. Journal Article 2018 Allegory of a cave crustacean: systematic and biogeographic reality of Halosbaena (Peracarida: Thermosbaenacea) sought with molecular data at multiple scales
Halse, S. Book Section 2018 Subterranean fauna of the arid zone
Page, T.J.,Stevens, M.I., Adams, M., Foster, R. Velasco-Castrillón, A. and Humphreys, W.F. Journal Article 2018 Multiple molecular markers reinforce the systematic framework of unique Australian cave fishes (Milyeringa : Gobioidei)
Perina, G., Camacho, A.I., Huey, J., Horwitz, P., and Koenders, A. Journal Article 2018 Understanding subterranean variability: the first genus of Bathynellidae (Bathynellacea, Crustacea) from Western Australia described through a morphological and multigene approach
Perina, G., Camacho, A.I., Huey, J., Horwitz, P. and Koenders, A. Journal Article 2019 New Bathynellidae (Crustacea) taxa and their relationships in the Fortescue catchment aquifers of the Pilbara region, Western Australia, Systematics and Biodiversity
Perina, G., Camacho, A., Huey, J., Horwitz, P. and Koenders, A. Journal Article 2019 The role of allopatric speciation and ancient origins of Bathynellidae (Crustacea) in the Pilbara (Western Australia): two new genera from the De Grey River catchment
Matthews, E.F., Abrams, K.M., Cooper, S.J.B., Huey, J.A., Hillyer, M.J., Humphreys, W.F., Austin, A.D. and Guzik, M.T. Journal Article 2019 Scratching the surface of subterranean biodiversity: molecular analysis reveals a diverse and previously unknown fauna of Parabathynellidae (Crustacea: Bathynellacea) from the Pilbara, Western Australia
Perina, G. Thesis 2019 Ancient origins of a diverse stygofaunal group: The Australian Bathynellidae (Crustacea)
White, N.E., Guzik, M.T., Austin, A.D., Moore, G.I. Humphreys, W.F., Alexander, J. and Bunce, M. Journal Article 2020 Detection of rare Australian endemic blind cave eel (Ophisternon candidum) with environmental DNA: implications for threatened species management in subterranean environments
Shelley, J.J., Swearer, S.E., Dempster, T., Adams, M., Le Feuvre, M.C., Hammer, M.P. and Unmack, P.J. Journal Article 2020 Plio-Pleistocene sea-level changes drive speciation of freshwater fishes in north-western Australia
Lintermans, M., Geyle, H.M., Beatty, S., Brown, C., Ebner, B.C., Freeman, R., Hammer, M.P., Humphreys, W.F., Kennard, M.J., Kern, P., Martin, K., Morgan, D.L., Raadik, T.A., Unmack, P.J., Wager, R., Woinarski, J.C.Z. and Garnett, S.T. Journal Article 2020 Big trouble for little fish: identifying Australian freshwater fishes in imminent risk of extinction
Eberhard, S.M. and Howarth, F.G. Journal Article 2021 Undara lava cave fauna in tropical Queensland with an annotated list of Australian subterranean biodiversity hotspots
Gierl, C., Dohrmann, M., Keith, P., Humphreys, W.J., Esmaeili, H.R., Vukić, J., Radek, S. and ReichenbacherI, B. Journal Article 2022 An integrative phylogenetic approach for inferring relationships of fossil gobioids (Teleostei: Gobiiformes)
Saccò, M., Guzik, M.T., van der Heyde, M., Nevill, P., Cooper, S.J.B., Austin, A.D., Coates, P.J. and Allentoft, M.E. Journal Article 2022 eDNA in subterranean ecosystems: Applications, technical aspects, and future prospects
Saccò, M., Blyth, A.J., Douglas, G., Humphreys, W.F., Hose, G.C., Davis, J., Guzik, M.T., Martínez, A., Eberhard, S.M. and Halse, S.A. Journal Article 2022 Stygofaunal diversity and ecological sustainability of coastal groundwater ecosystems in a changing climate: The Australian paradigm
Thacker, C.E., McCraney, W.T., Harrington, R.C., Near, T.J., Shelley, J.J., Adams, M., Hammer, M.P. and Unmack, P.J. Journal Article 2023 Diversification of the sleepers (Gobiiformes: Gobioidei: Eleotridae) and evolution of the root gobioid families
Webb, J.A., White, S.Q. and Smith, G.K. Book 2023 Australian caves and karst systems
White, S.Q., Brooks, D. and Webb, J.A. Book Section 2023 Cape Range
van der Heyde, M., Alexander, J., Nevill, P., Austin, A.D., Stevens, N., Jones, M. and Guzik, M.T. Journal Article 2023 Rapid detection of subterranean fauna from passive sampling of groundwater eDNA
van der Heyde, M., White, N.E., Nevill, P., Austin, A.D., Stevens, N., Jones, M. and Guzik, M.T. Journal Article 2023 Taking eDNA underground: factors affecting eDNA detection of subterranean fauna in groundwater