CURRENT PYRAMIDAL LIST OF
FAMILY-GROUP NAMES (FGN)
WITH INCLUDED (SUB)GENERA

 

SYNTHESIS | FGN LIST | FGN FOOTNOTES | FGN VS. TREE OF LIFE  |

 

 

This list of all Killifish family-group names and their included generic names is the most accurate available list, according to current research.

Synthesis

Killifish, the herein aggregated common name for all, oviparous and viviparous cyprinodontiformes fishes, are currently classified into 2 suborders (Aplocheiloidei, i.e., rivulins with hereafter first-listed 3 families, and Cyprinodontoidei, killifish in a strict sense, mostly, but also, pupfishes, toothcarps, lampeyes and livebearers, including splitfins) and in 14 families (full details HERE, constantly updated):
- Aplocheilidae, oviparous, only from India and Madagascar;
- Nothobranchiidae, oviparous, from Africa except Madagascar with 2 subfamilies: Nothobranchiinae and Epiplatinae;
- Rivulidae, oviparous, from Central and South America with 3 subfamilies: Rivulinae, Cynolebiinae and Kryptolebiatinae;
- Anablepsidae, with 3 subfamilies: Anablepsinae, viviparous, the monotypic Oxyzygonectinae, oviparous, and Jenynsiinae, viviparous, from Central and South America;
- Aphaniidae, oviparous, monotypic, from Europe, the Middle East, up to Pakistan and northwestern coastal India;
- Cyprinodontidae, oviparous, with 2 subfamilies: Cubanichthyinae (with 2 tribes, Cubanichthyini, Yssolebiini) and Cyprinodontinae (with 2 tribes, Cyprinodontini and Orestiadini), from North, Central and South America;
- Fluviphylacidae, oviparous, monotypic, from South America.
- Fundulidae, oviparous, from North and Central America;
- Goodeidae, with 2 subfamilies: Empetrichthyinae, oviparous and Goodeinae, viviparous, from North and Central America;
- Pantanodontidae, oviparous, previously monotypic, today with 4 extant genera, from Eastern Africa including Madagascar;
- Poeciliidae, with 3 subfamilies, from North, Central and South America: Poeciliinae viviparous, Tomeurinae, oviparous, monotypic, Xenodexiinae, viviparous, monotypic);
- Procatopodidae, oviparous, from Africa (in 2 subfamilies, Aplocheilichthyinae, monotypic, and Procatopodinae with 2 tribes, Procatopodini and Micropanchini);
- Profundulidae, oviparous, bitypic, from Central America;
- Valenciidae, oviparous, monotypic, from Europe;
These 14 families are also accepted in active international databases on all fishes to ensure coherence and universality, even if the major move from 7 to 14 families is very recent (2018) and is more reflecting the splitting trends in present systematics than a parsimonious analysis (a solid publication by Piller et al. in 2022 even goes to 16 families, likewise another solid and global phylogeny by Morales et al., 2024). The contents of lower levels depend more on authors : a consensus and conservative view is selected herein, fully in line with ICZN recommendations. Future general trend is probably moving to even more divisions at all family-group names levels, even if the pyramid may balance back to fewer families and consequently more lower levels in an un-foreseeable future (notably the first, but preliminary, analysis, by Teimori & Motamedi, in 2019, based on full mitogenome, discloses that family Aphaniidae is probably ambiguous and recent results tend to amalgamate Valenciidae into Aphaniidae).

Detailed list by alphabetical order (except for nomino-typical subunits, listed first, and synonyms, listed by historical order), based on stems (roots) of generic names, following I.C.Z.N. code main rule for family-group names (alternative spellings published by various authors are given after '#' following other rules of the same I.C.Z.N. code)

LAST UPDATE: June 26. 2024

Important notes : (1) since edition dated August 28. 2019, viviparous taxa are included (therefore present listing is 100% complete for Cyprinodontiformes), (2) Rivulidae are herein considered as maintained by ICZN but 2 contesting appeals by 2 independent lepidopterists are published late 2019 and ICZN commissioners will have to rule a decision (probably in 2022), (3) request to ICZN of homogenized changes for family-group names based on genera containing 'lebias', as -lebi, neither as -lebiat, nor as -lebias, except Kryptolebias is provisionally accepted, (4) Procatopodinae+Aplocheilichthyinae components are herein maintained conservatively, but still not fixed, pending an announced detailed publication by Bragança (in 2022), (5) lately Piller et al. have published a new molecular phylogeny of Cyprinodontiformes using a targeted approach, anchored hybrid enrichment, and their results increase total families to 16 (not 14) by upgrading Cubanichthyinae and Orestiadinae to full family rank (Morales et al., 2024, reach a similar conclusion of 16 families, based on 12 molecular markers)… those findings will be included herein as soon as international databases follow them (or not) [ref.: Piller, K.R., Parker, E., Lemmon, A.R. & Lemmon, E.M. 2022. Investigating the Utility of Anchored Hybrid Enrichment data to investigate the Relationships among the Killifishes (Actinopterygii: Cyprinodontiformes), a globally distributed Group of fishes. Molecular Biology and Evolution, https://doi.org/10.1016/j.ympev.2022.107482 ; Morales, P., F. Gajardo, C. Valdivieso, M. Valladares & M.L. Allende. 2024. Genomes of the Orestias pupfish from the Andean Altiplano shed Light on their evolutionary History and phylogenetic Relationships within Cyprinodontiformes. BMC Genomics, 25 (614). https://doi.org/10.1186/s12864-024-10416-w].

Beware, the automatic translator reverses scientific names : please use the English version to save this list.

Main reference on oviparous groups : Huber, J.H. 2005. A Review of Family-group names for oviparous Cyprinodontiformes (Pisces; Teleostei). British Killifish Ass. Publ., Separatum, (October): 16 pp., 1 tab. [also duplicated in Huber, J.H. 2006. Killi-Data Catalogue. Killi-Data Editions, Paris, (December 10.): 1028 pp., figs, maps, and freely available herein at Huber's dedicated PUBLICATION]
That publication is herein enriched with the bibliographic references of the family-group names and is systematically and nomenclaturally updated on a regular basis with the update date mentioned above, for new generic and existing taxa with footnotes for explanations and alternative options.
Main reference on viviparous groups : Huber, J.H. 2019. A nomenclatural and systematic Analysis of livebearing Cyprinodontiformes (Acanthopterygii: Anablepsinae, Goodeinae, Poeciliidae). Killi-Data Series 2019, 4-155, 3 tabs., 8 figs. [abstract : HERE].
Notes : according to I.C.Z.N. (ICZN) rules, the biological suprageneric pyramid is normally built based on the Latin or Greek root of the generic name with a suffix added in several levels, [1] the order (suffix -formes) basally {killifish are an historical artificial assemblage -initially many oviparous and rare viviparous species- of the order Cyprinodontiformes, today, herein, killifish and Cyprinodontiformes are identical}, [2] the sub-order (suffix -oidei), [3] the superfamily (suffix -oidea) {with limited usage today, due to very unstable evidence and conflicting results}, [4] the family (suffix -idae), [5] the subfamily (suffix -inae), [6] the tribe (suffix -ini), [7] the subtribe (suffix -ina), and at the bottom-end (before species names), the genus/subgenus levels ; the validity status and ranking of each generic taxon in the following list corresponds to the present evidence in Killi-Data ; must-read important footnotes highlight explanations, comments and/or clarifications in case of selective choices for the present update of the current pyramidal list of family-group names with included generic names.

Supra-order classification : Cyprinodontiformes are closely related to Beloniformes and Atheriniformes (most species from marine coastal fringes unlike killifishes), all part of superorder Atherinomorphae Greenwood, Rosen, Weitzman & Myers, 1966 (ray-finned fish) [Ref. : Greenwood, P.H., D.E. Rosen, S.H. Weitzman & G.S. Myers. 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bulletin of the American Museum of Natural History 131 (4): 339-456], as exemplified below [artificial pyramid, with no phylogeny relationship and no dating]. However, in September 2021, a new review [Ref. : Dornburg, A. & T.J. Near, 2021. The emerging phylogenetic Perspective on the Evolution of Actinopterygian Fishes. The Annual Review of Ecology, Evolution, and Systematics; 52: 427-452] proposes to lump Cyprinodontiformes and related orders into Blenniiformes, and if accepted everything in following tree and list will be downgraded by 2 steps and Cyprinodontiformes will become a superfamily only in suborder Atherinoidei, as Cyprinodontoidea… time will tell.

 

Full list of Cyprinodontiformes Family-Group Names 

 

Order Cyprinodontiformes Berg, 1940.
[Berg, L.S. 1940. The classification of fishes, both Recent and fossil. Trudy Zoologicheskogo lnstituta Akademiia nauk SSSR 5: 87-517]

 

Suborder Aplocheiloidei Bleeker, 1859.
[Bleeker, P. 1859. Enumeratio Specierum Piscium hucusque in Archipelage Indico Observatorum, adjectis Habitationibus Cattionibusque, ubi Descritiones earum recentiorus reperientur, nec non Specibus Musei Bleekeriani, bengalensibus, japonicis, capensibus, tasmanicisque. Act. Soc. Sc. Indo-Neerl., 6: i-xxxvi + 1-276. {page 30}]
- Synonym : Haplochileidei Garman, 1895 [emmendation at subfamily level, as Haplochilinae]
[Garman, S. 1895. The cyprinodonts. Memoirs of the Museum of Comparative Zoology at Harvard College 19 (1): 1-179, 12 plates. {page 93}]

 

Family Aplocheilidae Bleeker, 1859.

 

* Genus Aplocheilus McClelland, 1839.
- Synonym : Odontopsis Hasselt, 1823 (nomen oblitum)
- Synonym : Haplochilus Agassiz, 1846 (emendation)
- Synonym : Panchax Valenciennes in Cuvier & Valenciennes, 1846
- Unavailable : Homalopsis Kuhl & Hasselt in Bleeker, 1852

* Genus Pachypanchax Myers, 1933.

 

 

Family Nothobranchiidae Garman, 1895.
[Garman, S. 1895. The cyprinodonts. Memoirs of the Museum of Comparative Zoology at Harvard College 19 (1): 1-179, 12 plates. {page 159}]

 

Subfamily Nothobranchiinae Garman, 1895.

 

Tribe Nothobranchiini Garman, 1895.

 

Subtribe Nothobranchiina Garman, 1895.

 

* Genus Nothobranchius Peters, 1868.

* Subgenus Nothobranchius s.s. Peters, 1868.

* Subgenus Adiniops Myers, 1924.

* Subgenus Aphyobranchius Wildekamp, 1977.

* Subgenus Cynobranchius Costa, 2018.

* Subgenus Plesiobranchius Costa, 2018.

* Subgenus Paranothobranchius Seegers, 1985.

* Subgenus Zononothobranchius Radda, 1969.

* Genus Fundulosoma Ahl, 1924.

* Genus Pronothobranchius Radda, 1969.

 

Subtribe Aphyosemina Huber, 2000. (# name based on correct stem is Aphyosemiina, but Aphyosemiina may be seen unjustified following latest ICZN code for names created after 2000) (Footnotes 17 and 19)
[Huber, J.H. 2000. Killi-Data 2000. Updated checklist of taxonomic names, collecting localities and bibliographic references of oviparous Cyprinodont fishes (Cyprinodontiformes). Cybium, Soc. fr. Ichtyologie, Ed., Paris.: 538 pp., figs. {pages 28, 33}]

 

* Genus Aphyosemion Myers, 1924.

* Subgenus Aphyosemion s.s. Myers, 1924.

* Subgenus Chromaphyosemion Radda, 1971.

* Subgenus Diapteron Huber & Seegers, 1977.

* Subgenus Episemion Radda & Pürzl, 1987.

* Subgenus Iconisemion Huber, 2013.

* Subgenus Kathetys Huber, 1977.

* Subgenus Mesoaphyosemion Radda, 1977.

* Subgenus Raddaella Huber, 1977.

* Subgenus Scheelsemion Huber, 2013.

* Genus Foerschichthys Scheel & Romand, 1981.

* Genus Fundulopanchax Myers, 1924.

* Subgenus Fundulopanchax s.s. Myers, 1924.

* Subgenus Gularopanchax Radda, 1977.

* Subgenus Paludopanchax Radda, 1977.

* Subgenus Paraphyosemion Radda, 1977.

* Subgenus Pauciradius Wildekamp & Zee, 2005.

 

Tribe Adamansini Huber, 2000. (Footnote 6)
[Huber, J.H. 2000. Killi-Data 2000. Updated checklist of taxonomic names, collecting localities and bibliographic references of oviparous Cyprinodont fishes (Cyprinodontiformes). Cybium, Soc. fr. Ichtyologie, Ed., Paris.: 538 pp., figs. {pages 28, 30}]
- Synonym : Fenerbahceini Sonnenberg & Zee, 2008 (Footnote 6)
[Sonnenberg, R. & J.R. Van der Zee, 2008). On the validity of Fenerbahce Özdikmen et al., 2006 as replacement for Adamas Huber, 1979 (Cyprinodontiformes: Nothobranchiidae). Zootaxa, 1687, 67–68. {page 67}]

 

* Genus Fenerbahce Özdikmen, Polat, Yylmaz & Yazycyodlu, 2006. (Footnote 6)
- Synonym : Adamas Huber, 1979 (preoccupied)
- Synonym : Adamans Huber, 2007

 

Subfamily Epiplatinae Huber, 2000. (# name based on correct stem is Epiplateinae, but preoccupied) (Footnotes 7 and 17)
[Huber, J.H. 2000. Killi-Data 2000. Updated checklist of taxonomic names, collecting localities and bibliographic references of oviparous Cyprinodont fishes (Cyprinodontiformes). Cybium, Soc. fr. Ichtyologie, Ed., Paris.: 538 pp., figs. {pages 28, 32}]

 

 

Tribe Epiplatini Huber, 2000. (Footnote 7)

 

* Genus Epiplatys Gill, 1862.

* Subgenus Epiplatys s.s. Gill, 1862.

* Subgenus Aphyoplatys Clausen, 1967.

* Subgenus Kulonplatys Huber, 2024.

* Subgenus Lycocyprinus Peters, 1868.

* Subgenus Parepiplatys Clausen, 1967.

* Subgenus Pseudepiplatys Clausen, 1967.

* Subgenus Xerosplatys Huber, 2024.

 

Tribe Callopanchini Huber, 2000. (# name as Callopanchacini based on un-necessary stem) (Footnotes 8 and 19)
[Huber, J.H. 2000. Killi-Data 2000. Updated checklist of taxonomic names, collecting localities and bibliographic references of oviparous Cyprinodont fishes (Cyprinodontiformes). Cybium, Soc. fr. Ichtyologie, Ed., Paris.: 538 pp., figs. {pages 28, 31}]

 

* Genus Callopanchax Myers, 1933.
- Synonym : Roloffia Clausen, 1966 (invalid)

* Genus Archiaphyosemion Radda, 1977.

* Subgenus Archiaphyosemion s.s. Radda, 1977.

* Subgenus Nimbapanchax Sonnenberg & Busch, 2009.

* Genus Scriptaphyosemion Radda & Pürzl, 1987.

 

 

Family Rivulidae Myers, 1925. (Footnote 15)
[Myers, G.S. 1925. Results of some recent studies on the American killifishes. The Fish Culturist 4 (8): 370-371. {page 371}]

 

Subfamily Rivulinae Myers, 1925. (Footnote 15)

 

Tribe Rivulini Myers, 1925. (Footnote 15)

 

* Genus Rivulus Poey, 1860.

* Subgenus Rivulus s.s. Poey, 1860.

* Subgenus Anablepsoides Huber, 1992.

* Subgenus Atlantirivulus Costa, 2008.

* Subgenus Benirivulus Costa, 2006. (Footnote 24)

* Subgenus Cynodonichthys Meek, 1904.

* Subgenus Laimosemion Huber, 1999.

* Subgenus Melanorivulus Costa, 2006.

* Subgenus Oditichthys Huber, 1999.

* Subgenus Owiyeye Costa, 2006.

* Subgenus Vomerivulus Fowler, 1944.

 

Tribe Prorivulini Costa, 2008. (Footnote 16)
[Costa, W.J.E.M. 2008. Catalog of aplocheiloid Killifishes of the World. Universidade Federal do Rio de Janeiro Ed., Rio de Janeiro, Brasil, 127 pp. {page 77}]

 

* Genus Prorivulus Costa, 2004.

 

Tribe Neofundulini Costa, 1990. (Footnote 18)
[Costa, W.J.E.M. 1990. Classificaçao e Distribuçao da familia Rivulidae (Cyprinodontiformes, Aplocheiloidei). Revta. Brasil Biol., 50 (1): 83-89, figs. {page 85}]

 

Subtribe Neofundulina Costa, 1990.

 

* Genus Neofundulus Myers, 1924.

* Genus Trigonectes Myers, 1925.
- Synonym : Rivulichthys Myers, 1927

 

Subtribe Aphyolebiina Costa, 1998. (# names as Aphyolebiatina and Aphyolebiasina are respectively based on incorrect and un-necessary stems) (Footnotes 8, 16 and 17)
[Costa, W.J.E.M. 1998. Phylogeny and classification of Rivulidae revisited: Origin and evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei), J. Comp. Biol., 3 (1): 33-92. {page 78}]
- Synonym : Moemina Costa, 1998. (Footnotes, 8, 16)
[Costa, W.J.E.M. 1998. Phylogeny and classification of Rivulidae revisited: Origin and evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei), J. Comp. Biol., 3 (1): 33-92. {page 78}]

 

* Genus Moema Costa, 1989. (Footnote 16)

* Subgenus Moema s.s. Costa, 1989. (Footnote 16).

* Subgenus Aphyolebias Costa, 1998. (Footnote 16).

* Genus Micromoema Costa, 1998.

* Genus Renova Thomerson & Taphom, 1995.

 

Subtribe Pterolebiina Costa, 1990. (# names as Pterolebiatina and Pterolebiasina are respectively based on incorrect and un-necessary stems) (Footnotes 8 and 17)
[Costa, W.J.E.M. 1990. Classificaçao e Distribuçao da familia Rivulidae (Cyprinodontiformes, Aplocheiloidei). Revta. Brasil Biol., 50 (1): 83-89, figs. {page 87}]

 

* Genus Pterolebias Garman, 1895.

* Genus Gnatholebias Costa, 1998.

 

Tribe Rachoviini Costa, 1990. (# Rachovini, erratum) (Footnote 18)
[Costa, W.J.E.M. 1990. Classificaçao e Distribuçao da familia Rivulidae (Cyprinodontiformes, Aplocheiloidei). Revta. Brasil Biol., 50 (1): 83-89, figs. {page 85}]

 

Subtribe Rachoviina Costa, 1990. (# Rachovina, erratum)
- Synonym : Terranatina Costa, 1990.
[Costa, W.J.E.M. 1990. Classificaçao e Distribuçao da familia Rivulidae (Cyprinodontiformes, Aplocheiloidei). Revta. Brasil Biol., 50 (1): 83-89, figs. {page 86}]

 

* Genus Rachovia Myers, 1927.

* Genus Austrofundulus Myers, 1932.

* Genus Llanolebias Hrbek & Taphorn, 2008.

* Genus Terranatos Taphorn & Thomerson, 1978.

 

Subtribe Millerichthyina Costa, 1998.
[Costa, W.J.E.M. 1998. Phylogeny and classification of Rivulidae revisited: Origin and evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei), J. Comp. Biol., 3 (1): 33-92. {page 79}]

 

* Genus Millerichthys Costa, 1995.

 

Subtribe Plesiolebiina Costa, 1990. (# names as Plesiolebiatina and Plesiolebiasina are respectively based on incorrect and un-necessary stems) (Footnotes 8 and 17)
[Costa, W.J.E.M. 1990. Classificaçao e Distribuçao da familia Rivulidae (Cyprinodontiformes, Aplocheiloidei). Revta. Brasil Biol., 50 (1): 83-89, figs. {page 86}]

 

* Genus Plesiolebias Costa, 1989.

* Genus Maratecoara Costa, 1995.

* Genus Papiliolebias Costa, 1998.

* Genus Pituna Costa, 1989.

* Genus Stenolebias Costa, 1995.

 

Subfamily Cynolebiinae Hoedeman, 1961. (# names as Cynolebiatinae and Cynolebiasinae are respectively based on incorrect and un-necessary stems) (Footnotes 8 and 17)
[Hoedeman, J.J. 1961. Studies on Cyprinodontiform Fishes. 10. On the probable Evolution of the frontal Scalation Patterns. Bulletin of Aquatic Biology (Bull. Aquatic Biol.), 2 (18): 82-92, 2 figs. {page 87}]

 

Tribe Cynolebiini Hoedeman, 1961. (# names as Cynolebiatini and Cynolebiasini are respectively based on incorrect and un-necessary stems) (Footnotes 8 and 17)

 

Subtribe Cynolebiina Hoedeman, 1961. (# names as Cynolebiatina and Cynolebiasina are respectively based on incorrect and un-necessary stems) (Footnotes 8 and 17)

 

* Genus Cynolebias Steindachner, 1876.

* Subgenus Cynolebias s.s. Steindachner, 1876.

* Subgenus Bathylebias Costa, 2008.

* Genus Acantholebias Costa, 2008.

* Genus Acrolebias Costa, 2008.

* Genus Amatolebias Alonso, Teran, Serra, Calviño, Montes, García, Barneche, Almiron, Ciotek, Giorgis & Casciotta, 2023.

* Genus Argolebias Costa, 2008.

* Genus Austrolebias Costa, 1998.

* Genus Cypholebias Costa, 2008.

* Genus Garcialebias Alonso, Teran, Serra, Calviño, Montes, García, Barneche, Almiron, Ciotek, Giorgis & Casciotta, 2023.

* Genus Gymnolebias Costa, 2008.

* Genus Matilebias Alonso, Teran, Serra, Calviño, Montes, García, Barneche, Almiron, Ciotek, Giorgis & Casciotta, 2023.

* Genus Megalebias Costa, 1998.

* Genus Titanolebias Alonso, Teran, Serra, Calviño, Montes, García, Barneche, Almiron, Ciotek, Giorgis & Casciotta, 2023.

 

Spectrolebiina Costa, 1998. (# names as Spectrolebiatina and Spectrolebiasina are respectively based on incorrect and un-necessary stems) (Footnotes 8 and 17)
[Costa, W.J.E.M. 1998. Phylogeny and classification of Rivulidae revisited: Origin and evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei), J. Comp. Biol., 3 (1): 33-92. {page 74}]
- Synonym : Simpsonichthyina Costa, 1998 (Footnote 8).
[Costa, W.J.E.M. 1998. Phylogeny and classification of Rivulidae revisited: Origin and evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei), J. Comp. Biol., 3 (1): 33-92. {page 75}]

 

* Genus Simpsonichthys Carvalho, 1959.

* Subgenus Simpsonichthys s.s. Carvalho, 1959.

* Subgenus Hypsolebias Costa, 2006.

* Subgenus Ophthalmolebias Costa, 2006.

* Subgenus Spectrolebias Costa & Nielsen, 1997.

* Subgenus Xenurolebias Costa, 2006.

* Genus Nematolebias Costa, 1998.

 

Tribe Cynopoecilini Costa, 1990.
[Costa, W.J.E.M. 1990. Classificaçao e Distribuçao da familia Rivulidae (Cyprinodontiformes, Aplocheiloidei). Revta. Brasil Biol., 50 (1): 83-89, figs. {page 88}]

 

Subtribe Cynopoecilina Costa, 1990.

 

* Genus Cynopoecilus Regan, 1912.

* Subgenus Cynopoecilus s.s. Regan, 1912.

* Subgenus Poecilopanchax Costa, 2016.

* Genus Campellolebias Vaz-Ferreira & Sierra, 1974.

 

Subtribe Leptolebiina Costa, 1998. (# names as Leptolebiatina and Leptolebiasina are respectively based on incorrect and un-necessary stems) (Footnotes 8 and 17)
[Costa, W.J.E.M. 1998, Phylogeny and classification of Rivulidae revisited: Origin and evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei), J. Comp. Biol., 3 (1): 33-92. {page 73}]

 

* Genus Leptolebias Myers, 1952.

* Genus Leptopanchax Costa, 2016.

* Genus Mucurilebias Costa, 2014.

* Genus Notholebias Costa, 2008.

 

Subfamily Kryptolebiatinae Costa, 2004. (# name based on correct stem is Kryptolebiinae, but Kryptolebiinae is unjustified following latest ICZN code for names created after 2000) (Footnotes 8 and 17)
[Costa, W.J.E.M. 2004. Kryptolebias, a Substitute Name for Cryptolebias Costa, 2004 and Kryptolebiatinae, a Substitute Name for Cryptolebiatinae Costa, 2004 (Cyprinodontiformes: Rivulidae). Neotropical Ichthyology, 2 (2): 107-108. {page 107}]
- Nom. preoc. : Cryptolebiatinae Costa, 2004. (# name based on correct stem is Cryptolebiinae, but Cryptolebiinae is unjustified following latest ICZN code for names created after 2000) (Footnotes 8 and 17)
[Costa, W.J.E.M. 2004. Relationships and Redescription of Fundulus brasiliensis (Cyprinodontiformes: Rivulidae), with Description of a new Genus and Notes on the Classification of the Aplocheiloidei. Ichthyol. ExpIor. Freshwaters, 15 (2), 105-120, 10 figs., 3 tabs. {page 115}]

 

* Genus Kryptolebias Costa, 2004.
- Synonym : Cryptolebias Costa, 2004 (preoccupied)

 

 

 

Suborder Cyprinodontoidei Wagner, 1828.
[Wagner, R. 1828. Beiträge zur Kenntniss der Gattung Lebias Cuvier und der verwandten Gattungen, nebst Beschreibung zweier neuen in Sardinien entdeckten Arten. Isis (Oken), 21: 1050-1057. {page 1054]

 

Superfamily Cyprinodontoidea Wagner, 1828.

 

Family Cyprinodontidae Wagner, 1828.

 

Subfamily Cyprinodontinae Wagner, 1828.

 

Tribe Cyprinodontini Wagner, 1828.

 

* Genus Cyprinodon Lacepède, 1803.
- Synonym : Prinodon Rafinesque, 1815 (emendation)
- Synonym : Lebias Cuvier, 1816 (invalid)
- Synonym : Encrates Gistel, 1848
- Synonym : Trifarcius Poey, 1860

* Genus Cualac Miller, 1956.

* Genus Floridichthys Hubbs, 1926.

* Genus Garmanella Hubbs, 1936.

* Genus Jordanella Goode & Bean, 1879.

* Genus Megupsilon Miller & Walters, 1972.

 

Tribe Orestiadini Bleeker, 1859. (#Orestiasini #Orestiini #Orestiatini) (Footnote 12)
[Bleeker, P. 1859. Enumeratio Specierum Piscium hucusque in Archipelage Indico Observatorum, adjectis Habitationibus Cattionibusque, ubi Descritiones earum recentiorus reperientur, nec non Specibus Musei Bleekeriani, bengalensibus, japonicis, capensibus, tasmanicisque. Act. Soc. Sc. Indo-Neerl., 6: i-xxxvi + 1-276. {page 30}]

 

* Genus Orestias Valenciennes, 1839.

* Genus Pseudorestias Arratia, Vila, Lam, Guerrero & Quezada, 2017.
- Unavailable : Protorestias Eigenmann & Allen, 1942

 

Subfamily Cubanichthyinae Parenti, 1981.
[Parenti, L.R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History 168 (4): 335-557, 99 figs., 3 tabs, maps. {page 519}]

 

Tribe Cubanichthyini Parenti, 1981

* Genus Cubanichthys Hubbs, 1926.

* Genus Chriopeoides Fowler, 1939. (Footnote 5)

 

Tribe Yssolebiini Huber, 2015
[Huber, J.H. 2015. A morphological Rediagnosis of Yssolebias within cyprinodontoids (Cyprinodontiformes) following the detailed osteological Analysis by Costa based on a new Radiograph of the single Type of Cyprinodon martae Steindachner. Killi-Data Series 2015, 4-16, 3 figs., 2 tabs. {page 13}]

* Genus Yssolebias Huber, 2012. (Footnote 14)

 

 

Family Anablepsidae Bonaparte, 1831. (#Anablepidae #Anableptidae) [viviparous, in part] (Footnote 13)
[Bonaparte, C.L. Princ. 1831. Animali vertebrati. Prospetto del Sistema Generale d'Ittiologia, Tavola Metodica. Roma. Presso Antonio Boulzaler, vol. 52, 89-123; 155-189. {page 160, 179}]

 

Subfamily Anablepsinae Bonaparte, 1831. (#Anablepinae #Anableptinae) [viviparous] (Footnote 13)

 

Tribe Anablepsini Bonaparte, 1831. (#Anablepinae #Anableptinae) [viviparous] (Footnote 13)

 

* Genus Anableps Scopoli, 1877.
- Synonym : Peltatetraops Myers & Fowler, 1931

 

Tribe Jenynsiini Günther, 1866. [viviparous]
[Günther, A. 1866. Catalog of Fishes in the British Museum. Catalogue of the Physostomi, containing the families Salmonidae, Percopsidae, Galaxidae, Mormyridae, Gymnarchidae, Esocidae, Umbridae, Scombresocidae, Cyprinodontidae, in the collection of the British Museum. British Museum, London, 6: i-xv +368 pp. {page 300}]
- Synonym : Fitzroyiini Henn, 1916
[Henn, A.W. 1916. On various South American Poeciliid Fishes. Ann. Carnegie Mus., 10 (9): 93-142, pls. 18-21. {page 96}]

 

* Genus Jenynsia Günther, 1866.

* Subgenus Jenynsia s.s. Günther, 1866.
- Synonym : Fitzroyia Günther, 1866

* Subgenus Plesiojenynsia Ghedotti, 1998.

 

Subfamily Oxyzygonectinae Parenti, 1981. [oviparous]
[Parenti, L.R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History 168 (4): 335-557, 99 figs., 3 tabs, maps. {page 504}]

 

* Genus Oxyzygonectes Fowler, 1916.

 

 

Family Aphaniidae Hoedeman, 1949. (Footnote 23)
[Hoedeman, J.J. 1949. Family Cyprinodontidae, In: Hoedeman, J.J. & J.C.M. de Jong, Eds. 1947 - 1958. Encyclopaedie voor den aquariumhouder, De Regenboog, Amsterdam, The Netherlands, looseleaf edition, 56 parts. {X, 40-21, page 5}]
- Nom. oblitum : Telliini Bleeker, 1864. (Footnote 2)
[Bleeker, P. 1864. Atlas ichthyologique des Indes Orientales Néerlandaises, publié sous les Auspices du Gouvernement Néerlandais III. Cyprins. Frederic Muller, Amsterdam, (1863): 1-150, 43 pls. {page 139}] - Invalid name : Lebiatina Costa, 1997. (# name based on correct stem is Lebiina) (based on Lebias Goldfuss, 1820, a rejected name following ICZN Opinion 2057)
[Costa, W.J.E.M. 1997. Phylogeny and Classification of the Cyprinodontidae revisited (Teleostei: Cyprinodontiformes), are Andean and Anatolian killifishes sister taxa? Journal of Comparative Biology, 2 (1), 1-17. {page 13}]

 

* Genus Aphanius Nardo, 1827.

* Subgenus Aphanius s.s. Nardo, 1827.
- Synonym : Micromugil Gulia, 1861

* Subgenus Apricaphanius Freyhof & Yogurtçuoglu, 2020. (Footnote 23)

* Subgenus Anatolichthys Kosswig & Sözer, 1945.
- Synonym : Turkichthys Ermin, 1946

* Subgenus Esmaeilius Freyhof & Yogurtçuoglu, 2020.

* Subgenus Kosswigichthys Sözer, 1942.

* Subgenus Tellia Gervais, 1853.

* Genus Paraphanius Esmaeili, Teimori, Zarei & Sayyadzadeh, 2020.

* Genus Aphaniops Hoedeman, 1951.

 

 

Family Fundulidae Günther, 1866. (Footnote 1)
[Günther, A. 1866. Catalog of Fishes in the British Museum. British Museum, London, 6: 368 pp. {page 299}]
- Nom. oblitum : Hydrargiridae Gill, 1861 (Footnote 2)
[Gill, T. 1861. Catalog of the Fishes of the eastern Coast of North America from Greenland to Georgia. Proc. Acad. Nat. Sci. Philad., 13 (1860) (supplement): 1-63. {page 51}]

 

* Genus Fundulus Lacepède, 1803. (Footnote 9)

* Subgenus Fundulus s.s. Lacepède, 1803.
- Synonym : Hydrargira Lacepède, 1803
- Synonym : Borborys Goode & Bean, 1885

* Subgenus Fontinus Jordan & Evermann, 1896.
- Synonym : Galasaccus Fowler, 1916

* Subgenus Plancterus Garman, 1895.

* Subgenus Wileyichthys Ghedotti & Davis, 2013.

* Subgenus Xenisma Jordan in Jordan & Copeland, 1877.
- Synonym : Gambusinus Jordan & Evermann, 1896

* Subgenus Zygonectes Agassiz, 1854.
- Synonym : Adinia Girard, 1859
- Synonym : Micristius Gill, 1865

* Genus Leptolucania Myers, 1924.

* Genus Lucania Girard, 1859.
- Synonym : Chriopeops Fowler, 1916

 

 

Family Fluviphylacidae Roberts, 1970. (Footnote 20)
[Roberts, T.R. 1970. Description, osteology and relationships of the Amazonian cyprinodont fish Fluviphylax pygmaeus (Myers and Carvalho). Breviora (347): 28 pp., 13 figs. {page 22}]

 

* Genus Fluviphylax Whitley, 1965.
- Synonym : Potamophylax Myers & Carvalho in Myers, 1955 (preoccupied name)

 

 

Family Goodeidae Jordan & Gilbert, 1883. [viviparous, in part]
[Jordan, D.S. & C.H. Gilbert. 1883. A synopsis of the Fishes of North America. Bulletin of the United States National Museum 16: 1-1018. {page 327}]

 

Subfamily Goodeinae Jordan & Gilbert, 1883. [viviparous] (Footnote 21)

 

Tribe Goodeini Jordan & Gilbert, 1883.

 

Subtribe Goodeina Jordan & Gilbert, 1883.

 

* Genus Goodea Jordan, 1880.
- Synonym : Xenendum Jordan & Snyder, 1899.

 

Subtribe Ataeniobiina Hubbs & Turner, 1939.
[Hubbs, C.L. & C.L. Turner. 1939. Studies of the fishes of the order Cyprinodontes. XVI. A revision of the Goodeidae. Miscellaneous Publications of the Museum of Zoology University of Michigan (Misc. Publ. Mus. Zool. Univ. Michigan), 42 (9 Nov.): 1-80, 5 pls. {page 39}]

* Genus Ataeniobius Hubbs & Turner, 1939.

 

Tribe Zoogoneticini Hubbs, 1924.
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. I. A Classification of the fishes of the Order. II. An Analysis of the genera of the Poeciliidae. III. The species of Profundulus a new genus from Central America. IV.The subspecies of Pseudoxiphophorus bimaculatus and of Priapichthys annectens. Miscellaneous Publications of the Museum of Zoology University of Michigan, 13 (Jan. 18.): 31 pp., 4 pls., tabs. {page 4}]
- Synonym : Chapalichthyini Doadrio & Dominguez, 2004
[Doadrio, I. & O.D. Dominguez. 2004. Phylogenetic Relationships within the fish family Goodeidae based on Cytochrome b Sequence Data. Molecular Phylogenetics and Evolution (Mol. Phylogenet. Evol.), 31 (2) (May): 416-430. {page 424}]

* Genus Zoogoneticus Meek, 1902.

* Genus Alloophorus Hubbs & Turner, 1939.

* Genus Ameca Miller & Fitzsimons, 1971.

* Genus Chapalichthys Meek, 1902. (Footnote 21)

* Genus Xenoophorus Hubbs & Turner, 1939.

* Genus Xenotoca Hubbs & Turner, 1939. (Footnote 21)

 

Tribe Characodontini Regan, 1907.
[Regan, C.T. 1907. Pisces. In: Biologia Centrali Americana. P. Press, 8 (May): i-xxxiii + 1-203, 12 figs., 2 maps, 26 pls., tabs., keys. {page 76}]

 

* Genus Characodon Günther, 1866.

 

Tribe Girardinichthyini Hubbs & Turner.
[Hubbs, C.L. & C.L. Turner. 1939. Studies of the fishes of the order Cyprinodontes. XVI. A revision of the Goodeidae. Miscellaneous Publications of the Museum of Zoology University of Michigan (Misc. Publ. Mus. Zool. Univ. Michigan), 42 (9 Nov.): 1-80, 5 pls. {page 57}]

 

* Genus Girardinichthys Bleeker, 1860.
- Synonym : Limnurgus Günther, 1866.
- Synonym : Lermichthys Hubbs, 1926.
- Synonym : Hubbsina Buen, 1941. (Footnote 21)

* Genus Allotoca Hubbs & Turner, 1939.
- Synonym : Neoophorus Hubbs & Turner, 1939.

* Genus Neotoca Hubbs & Turner, 1939. (Footnote 21)

* Genus Skiffia Meek, 1902.
- Synonym : Ollentodon Hubbs & Turner, 1939.

 

Tribe Ilyodontini Doadrio & Dominguez, 2004.
[Doadrio, I. & O.D. Dominguez. 2004. Phylogenetic Relationships within the fish family Goodeidae based on Cytochrome b Sequence Data. Molecular Phylogenetics and Evolution (Mol. Phylogenet. Evol.), 31 (2) (May): 416-430. {page 424}]

 

* Genus Ilyodon Eigenmann, 1907.
- Synonym : Balsadichthys Hubbs, 1926.

* Genus Allodontichthys Hubbs & Turner, 1939.

* Genus Xenotaenia Turner, 1946.

 

Subfamily Empetrichthyinae Jordan, Evermann, & Clark, 1930. [oviparous]
[Jordan, D.S., B.W. Evermann & H.W. Clark. 1930. Checklist of the Fishes and Fish-like Vertebrates of North and Middle America north of the Northern boundary of Venezuela and Colombia. U.S. Comm. of Fish. Rept. for 1928, Part 2, App. 10: 670pp. {page 182}]

 

* Genus Empetrichthys Gilbert, 1893.

* Genus Crenichthys Hubbs, 1932.

 

 

Family Poeciliidae Bonaparte, 1831. [viviparous, in part] (Footnotes 11 and 20)
[Bonaparte, C.L. Princ. 1831. Animali vertebrati. Prospetto del Sistema Generale d'Ittiologia, Tavola Metodica. Roma. Presso Antonio Boulzaler, vol. 52, 89-123; 155-189. {page 123}]

 

Subfamily Poeciliinae Bonaparte, 1831. [viviparous]

 

Tribe Poeciliini Bonaparte, 1831.
- Synonym : Pamphoriini Hubbs, 1924.
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. Misc. Papers Mus. Zool. Univ. Michigan, 13: 31 pp., 4 pls., tabs. {page 10}]

 

* Genus Poecilia Bloch & Schneider, 1801. (Footnote 22)

* Subgenus Poecilia s.s. Bloch & Schneider, 1801.
- Synonym : Alazon Gistel, 1848.
- Synonym : Lebistes Filippi, 1861. (Footnote 22)
- Synonym : Neopoecilia Hubbs, 1924.

* Subgenus Acanthophacelus Eigenmann, 1907.

* Subgenus Allopoecilia Hubbs, 1924.
- Synonym : Hubbsichthys Schultz, 1949.

* Subgenus Curtipenis Rivas & Myers, 1950.

* Subgenus Limia Poey, 1854.
- Synonym : Acropoecilia Hilgendorf, 1889.
- Synonym : Odontolimia Rivas, 1980.

* Subgenus Micropoecilia Hubbs, 1926.
- Synonym : Recepoecilia Whitley, 1951.

* Subgenus Mollienesia LeSueur, 1821.
- Synonym : Lembesseia Fowler, 1949.

* Subgenus Pamphorichthys Regan, 1913.
- Nom. preoc. : Pamphoria Regan, 1913.
- Synonym : Parapoecilia Hubbs, 1924.

* Subgenus Pseudolimia Poeser, 2002.

* Subgenus Psychropoecilia Myers, 1935.

 

Tribe Belonesocini Bleeker, 1864. (Footnote 22)
[Bleeker, P. 1864. Atlas ichthyologique des Indes Orientales Néerlandaises, publié sous les Auspices du Gouvernement Néerlandais III. Cyprins. Frederic Muller, Amsterdam, (1863): 1-150, 43 pls. {page 140}]

 

Subtribe Belonesocina Bleeker, 1864.

 

* Genus Belonesox Kner, 1860.

 

Subtribe Gambusiina Gill, 1889.
[Gill, T.N. 1889. Articles. in: The Century Dictionary. An Encyclopedic Lexicon of the English Language. The Century Company, New York, 3: 2423-3556. {page 2446}]
- Synonym : Dicerophallini Alvarez, 1952.
[Alvarez, J.V. 1952. Dicerophallini, nueva Tribu de Poeciliidae de Chiapas (Pisc., Cyprinodont.). Ciencia (Patronato de Ciencia), 12 (3-4) (15 Aug.): 95-97, 3 figs., 1 tab. {page 95}]

 

* Genus Gambusia Poey, 1854.

* Subgenus Gambusia s.s. Poey, 1854.

* Subgenus Arthrophallus Hubbs, 1926.
- Synonym : Schizophallus Hubbs, 1926.

* Subgenus Heterophallina Hubbs, 1926.
- Synonym : Flexipenis Hubbs in Rivas, 1963.

* Subgenus Paragambusia Meek, 1904.
- Synonym : Orthophallus Rivas, 1963.
- Synonym : Toluichthys Dahl in Dahl & Medem, 1964.

* Genus Heterophallus Regan, 1914.
- Synonym : Dicerophallus Alvarez, 1952.

 

Tribe Cnesterodontini Hubbs, 1924. (Footnote 22)
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. Misc. Papers Mus. Zool. Univ. Michigan, 13: 31 pp., 4 pls., tabs. {page 9}]

 

* Genus Cnesterodon Garman, 1895.
- Synonym : Gulapinnus Langer, 1913.

* Genus Phalloceros Eigenmann, 1907.

* Genus Phalloptychus Eigenmann, 1907.

* Genus Phallotorynus Henn, 1916.

 

Tribe Girardinini Hubbs, 1924.
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. Misc. Papers Mus. Zool. Univ. Michigan, 13: 31 pp., 4 pls., tabs. {page 9}]

 

Subtribe Girardinina Hubbs, 1924.

 

* Genus Girardinus Poey, 1854.

* Subgenus Girardinus s.s. Poey, 1854.
- Synonym : Toxus Eigenmann, 1903.
- Synonym : Allodontium Howell-Rivero & Rivas, 1944.

* Subgenus Dactylophallus Howell-Rivero & Rivas, 1944.

* Subgenus Glaridichthys Garman, 1896.
- Nom. preoc. : Glaridodon Garman, 1895.

 

Subtribe Quintanina Howell-Rivero & Rivas, 1944.
[Howell-Rivero, L. & L.R. Rivas. 1944. Studies of cyprinodont fishes. Two new genera of the tribe Girardinini, from Cuba. Torreia (La Habana), 12 (May 31.): 1-19, 2 pls. {page 5}]

 

* Genus Quintana Hubbs, 1934.

 

Tribe Heterandriini Hubbs, 1924. (Footnote 22)
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. Misc. Papers Mus. Zool. Univ. Michigan, 13: 31 pp., 4 pls., tabs. {page 7}]

 

Subtribe Heterandriina Hubbs, 1924. (Footnote 22)

 

* Genus Heterandria Agassiz, 1853.

* Genus Pseudopoecilia Regan, 1913.

* Genus Diphyacantha Henn, 1916.
- Synonym : Darienichthys Hubbs, 1924.

* Genus Panamichthys Hubbs, 1924.

 

Subtribe Alfarina Hubbs, 1924.
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. Misc. Papers Mus. Zool. Univ. Michigan, 13: 31 pp., 4 pls., tabs. {page 11}]

 

* Genus Alfaro Meek, 1912.
- Nom. preoc. : Petalosoma Regan, 1908.
- Synonym : Petalurichthys Regan, 1912.
- Synonym : Furcipenis Hubbs, 1931.

* Genus Xenophallus Hubbs, 1924.

 

Subtribe Brachyrhaphina Lucinda & Reis, 2005. (Footnote 22)
[Lucinda, P.H.F. & T.E. Reis. 2005. Systematics of the subfamily Poeciliinae Bonaparte (Cyprinidontiformes: Poeciliidae), with an Emphasis on the tribe Cnesterodontini Hubbs. Neotrop. ichthyol., 3 (1) (31 Mar.): 1-60. {page 38}]

 

* Genus Brachyrhaphis Regan, 1913.
- Synonym : Plectrophallus Fowler, 1932.
- Synonym : Trigonophallus Hubbs, 1926.

* Genus Phallichthys Hubbs, 1924.

 

Subtribe Priapichthyina Lucinda & Reis, 2005. (Footnote 22)
[Lucinda, P.H.F. & T.E. Reis. 2005. Systematics of the subfamily Poeciliinae Bonaparte (Cyprinidontiformes: Poeciliidae), with an Emphasis on the tribe Cnesterodontini Hubbs. Neotrop. ichthyol., 3 (1) (31 Mar.): 1-60. {page 38}]

 

* Genus Priapichthys Regan, 1913.
- Synonym : Alloheterandria Hubbs, 1924.

* Genus Hiatirhaphis Huber, 2019.

 

Tribe Poeciliopsini Hubbs, 1924.
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. Misc. Papers Mus. Zool. Univ. Michigan, 13: 31 pp., 4 pls., tabs.]

 

* Genus Poeciliopsis Regan, 1913.

* Subgenus Poeciliopsis s.s. Regan, 1913.
- Invalid : Hemixiphophorus Bleeker, 1860.
- Synonym : Poecilistes Hubbs, 1926.

* Subgenus Aulophallus Hubbs, 1926.

* Subgenus Leptorhaphis Regan, 1913.
- Synonym : Arizonichthys Nichols, 1940.

* Genus Neoheterandria Henn, 1916.
- Synonym : Allogambusia Hubbs, 1924.

 

Tribe Priapellini Ghedotti, 2000.
[Ghedotti, M.J. 2000. Phylogenetic Analysis and Taxonomy of the Poecilioid Fishes (Teleostei: Cyprinodontiformes). Zool. J. Linnean Soc. London, 130 (1) (September), 1-53, 4 tabs., 18 figs. {page 39}]

 

* Genus Priapella Regan, 1913.

 

Tribe Scolichthyini Rosen, 1967. (Footnote 22)
[Rosen, D.E. 1967. New poeciliid fishes from Guatemala, with Comments on the Origins of some South and Central American Forms. Amer. Mus. Novitates, 2303 (Oct. 20.): 1-15. {page 2}]

 

* Genus Scolichthys Rosen, 1967.

* Genus Carlhubbsia Whitley, 1951.
- Nom. preoc. : Allophallus Hubbs, 1936.

 

Tribe Xiphophorini Hubbs, 1924.
[Hubbs, C.L. 1924. Studies of the Fishes of the Order Cyprinodontes. Misc. Papers Mus. Zool. Univ. Michigan, 13: 31 pp., 4 pls., tabs. {page 10}]

 

* Genus Xiphophorus Heckel, 1848.
- Synonym : Platypoecilus Günther, 1866.

* Genus Pseudoxiphophorus Bleeker, 1860. (Footnote 22)
- Synonym : Poeciliodes Steindachner, 1863.

 

Subfamily Tomeurinae Eigenmann, 1912. [oviparous] (Footnote 3)
[Eigenmann, C.H. 1912 The freshwater fishes of British Guiana, including a study of the ecological grouping of species, and the relation of the fauna of the plateau to that of the lowlands. Memoirs of the Carnegie Museum, 5 (1): i-xxii + 1-578p, pls. 1-103 {page 460}]

 

* Genus Tomeurus Eigenmann, 1909.

 

Subfamily Xenodexiinae Hubbs, 1950. [viviparous] (Footnote 3)
[Hubbs, C.L. 1950. Studies of cyprinodont Fishes. XX. A new subfamily from Guatemala, with ctenoid Scales and a unilateral pectoral Clasper. Misc. Publ. Mus. Zool. Univ. Michigan, 78 (Dec. 28.): 1-28, 4 pls. {page 5}]

 

* Genus Xenodexia Hubbs, 1950.

 

 

Family Profundulidae Hoedeman & Bronner, 1951. [oviparous] (Footnote 10)
[Hoedeman, J.J. & F.J. Bronner. 1951. De orde van de Tandkarpertjes, Cyprinodontiformes Berg, 1940. Deel VI. Het Aquarium (The Netherlands) 22 (1): 6-10, figs. {pages 7, 9}]

 

* Genus Profundulus Hubbs, 1924.

* Genus Tlaloc Alvarez & Carranza, 1951 (Footnote 10)

 

Family Procatopodidae Fowler, 1916. [oviparous] (# Procatopinae, erratum) (Footnote 20)
[Fowler, H.W. 1916. Notes on Fishes of the Orders Haplomi and Microcyprini. Proc. Acad. Nat. Sci. Philad., 68: 415-439. {page 416}]

 

Subfamily Procatopodinae Fowler, 1916. (# Procatopinae, erratum) (Footnote 20)

 

Tribe Procatopodini Fowler, 1916. (# Procatopini, erratum)
- Synonym : Lamprichthyina Fowler, 1916.

 

* Genus Procatopus Boulenger, 1904.
- Synonym : Andreasenius Clausen, 1959

* Genus Aapticheilichthys Huber, 2011.

* Genus Hylopanchax Poll & Lambert, 1965.

* Genus Hypsopanchax Myers, 1924.

* Genus Lamprichthys Regan, 1911.
- Synonym : Mohanga Boulenger, 1911

* Genus Plataplochilus Ahl, 1928.

* Genus Platypanchax Ahl, 1928.

* Genus Rhexipanchax Huber, 1999.

 

Tribe Micropanchini Huber, 2000.
[Huber, J.H. 2000. Killi-Data 2000. Updated checklist of taxonomic names, collecting localities and bibliographic references of oviparous Cyprinodont fishes (Cyprinodontiformes). Cybium, Soc. fr. Ichtyologie, Ed., Paris.: 538 pp., figs. {page 29, 55}]

 

* Genus Micropanchax Myers, 1924.

* Genus Congopanchax Poll, 1971.

* Genus Laciris Huber, 1981.

* Genus Lacustricola Myers, 1924.

* Subgenus Lacustricola s.s. Myers, 1924.

* Subgenus Cynopanchax Ahl, 1928.

* Genus Poropanchax Clausen, 1967.

 

Subfamily Aplocheilichthyinae Myers, 1928. (Footnote 20)
[Myers, G.S. 1928. The systematic Position of the Phallosthetid Fishes, with Diagnosis of a new Genus, from Siam. American Museum Novitates, n°295: 1-12. {page 4}]

 

* Genus Aplocheilichthys Bleeker, 1862.
- Synonym : Haplochilichthys Garman, 1895 (emendation)

 

 

Family Valenciidae Parenti, 1981. [oviparous] (Footnote 4)
[Parenti, L.R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History 168 (4): 335-557, 99 figs., 3 tabs, maps. {page 499}]

 

* Genus Valencia Myers, 1928.

 

 

Superfamily Pantanodontoidea Myers, 1955. [oviparous] (Footnote 20)
[Myers, G.S. 1955. Notes on the Classification and Names of Cyprinodont Fishes. Tropical Fish Magazine, 4 (4): 7. {page 7}]

 

Family Pantanodontidae Myers, 1955. (Footnote 20)

* Genus Pantanodon Myers, 1955. (Footnote 20)

* Genus Malagodon Meinema & Huber, 2023. (Footnote 20)

* Genus Aliteranodon Meinema & Huber, 2023. (Footnote 20)

* Genus Eremodon Huber & Meinema, 2024. (Footnote 20)

 

 

 

Important footnotes on Cyprinodontiformes Family-Group names 

 

Footnote 1 : Laan (pers. comm. to Huber, November 7. 2005) demonstrates Günther's priority over Jordan & Gilbert [Jordan, D.S. & C.H. Gilbert. 1883. A synopsis of the fishes of North America. Bulletin of the United States National Museum 16: 1-1018]

Footnote 2 : nomen oblitum is according to new concept in ICZN code (1999), valid from January 1. 2000 (following Huber, 2005).

Footnote 3 : Lucinda & Reis (2005), using osteology and morphology, place the genus Tomeurus as the most primitive taxon among Poeciliinae and in a separate tribe, whereas Ghedotti (2000), using the same type of characters (osteology and morphology) places it very derived and closest to the southern genus Cnesterodon, as a subtribe within Cnesterodontini ; Tomas Hrbek, Jens Seckinger and Axel Meyer (2007) or B.J.A. Pollux, R.W. Meredith, M.S. Springer, T. Garland and D.N. Reznick (2014) or Reznick D.N., A.I. Furness, R.W. Meredith & M.S. Springer. 2017, using molecular techniques, place Tomeurus, as the most primitive taxon of their matrix, except Xenodexia ctenolepis : since it is not the most primitive, the authors conclude, like Ghedotti (but for different reasons), that the live-bearing capacities were lost in genus Tomeurus ; these conflicting results are impossible to translate into a stable consensus picture and, according to latest published molecular evidences, the genus Tomeurus is herein positioned in a separate subfamily (not a tribe) in Poeciliidae, and, the genus Xenodexia would be positioned in another separate subfamily (not a tribe) in Poeciliidae, until better evidence.

Footnote 4 : according to preliminary recent fossil and molecular evidence, Valencia is more related to Aphanius, whereas according to recent osteological evidence (cranial soft tissue characters), Valencia is more related to Fundulidae, hence in both cases the family status Valenciidae might be discontinued in the future ; until stabilization, Valenciidae are maintained here, and this may be strengthened by a recent study of fossil fishes by Costa (2012) in which 2 extinct genera, Prolebias (in a revised restricted sense) and Francolebias are directly related to Valenciidae.

Footnote 5 : validity based on unpublished molecular evidence and variation of teeth structure according to Ghedotti (2000).

Footnote 6 : controversial case, with Huber (2007) analysing Fenerbahce as an electronic unavailable name replacing Adamas by Adamans, and Özdikmen (2008), Sonnenberg & Zee (2008), analysing Fenerbahce as available. Dubois (2008) demonstrates as valid the replacement family-group name Adamansini, according to ICZN code (valid from year 2000).

Footnote 7 : Epiplateinae moved back to Epiplatinae, according to ICZN preoccupied name (in Diptera), by Huber (2012) ; systematics of genera and subgenera are very unstable with no modern diagnoses ; according to extended molecular analysis by Collier (2016a) {but with still low bootsraps, no rediagnoses and opposite results from osteological evidence), Epiplatys is considered as a single genus in 2 clades, western and savannah clade and eastern and coastal clade, the latter hypothesized as having invaded inland following a Ogooué-Ivindo route {but the author discusses possibly also 4 valid genera, (1) Epiplatys s.s., with sexfasciatus-multifasciatus-grahami-sangmelinensis groups, possibly with several subgenera, one only being already named as Parepiplatys for grahami group, (2) monotypic Aphyoplatys, (3) un-named genus with fasciolatus-bifasciatus-spilargyreius groups, (4) Lycocyprinus, with dageti-chaperi groups, separated or not from Pseudepiplatys.

Footnote 8 : Costa creates many family-group names in 1998 including at levels not recognized by ICZN (e.g., infrafamily, supertribe) [resp. Leptolebiatini page 73, Spectrolebiatini page 74, Simpsonichthyina, page 75, Aphyolebiatina page 78 first line, Moemina page 78, 15 lines before end, Millerichthyini page 79], however it is important to note, following ICZN code (though it may seem strange and over-complicated) that Spectrolebiatina has priority over related Simpsonichthyina and the latter is today synonymized, even if generic name Spectrolebias is junior and a subgenus of generic name Simpsonichthys, and that Aphyolebiatina has priority over related Moemina and the latter is today synonymized, even if generic name Aphyolebias is junior and very related, a subgenus, maybe a junior synonym of generic name Moema ; besides, Costa (1998) uses all family-group names built from Lebias with the root lebiat- (e.g., Cynolebiatinae, Plesiolebiatini, Leptolebiatini) until 2008 when he moves to the root with the full name as lebias- (e.g., Cynolebiasinae, Plesiolebiasini, Leptolebiasini), disregarding Huber's previous revision (2005) ; Laan, Eschmeyer & Fricke (2014) have shown those are un-necessary changes, even if they fit, in part, with ICZN rules. Laan (2019) requests as a ruling by ICZN that all family-group names built from Lebias are built as lebi-, neither lebiat-, nor lebias-, (except Kryptolebiatinae, described after 2000) and it is herein followed as latest evidence, pending ICZN ruling if challenged [ref. Laan, R. van der. 2019. Case 3793- Request for a Confirmation of the correct Stem for Family-Group Names proposed before 2000 and typified by fish genera with names ending in -lebias. Bulletin of Zoological Nomenclature, 76: 48-52] ; Plesiolebiina are, depending on authors, either placed related to Rivulus et al. genera or to Cynolebias et al. genera, but latest molecular evidence favors first option, though with limited bootstrap support.

Footnote 9: the name is spelt either Lacépède or, as herein, Lacepède, following a detailed study by the senior, now deceased, French ichthyologist, Jacques Daget, at Paris MNHN… but it seems that French scientists could not reach a common conclusion after his death and both spellings are found everywhere on the Internet… when the issue is definitely sorted out, the correct spelling will be herein adopted.

Footnote 10: recent evidence by Meyer & Lydeard (1993), Webb et al. (2004), Resnick et al. (2017), using molecular techniques, show that Profundulus is not related to Fundulus et al., but to the viviparous Goodeinae (and this has been confirmed by Costa, 1998, on osteology) and to Crenichthys/Empetrichthys ; again, Ghedotti & Davis, 2013, combining morpho-osteology and molecular biology, tend to place Profundulidae closer to Goodeidae than to Fundulidae ; Tlaloc is only recently considered as a distinct genus from Profundulus based on molecular data and consensus of authors between 2016 and 2019 (previously as a simple junior synonym or conservatively as a subgenus).

Footnote 11: Laan (pers. comm. to Huber, March 4. 2013) reports that the family-group names derived from Anableps and from Poecilia have been described by Bonaparte on the same page (page 160, as Anableptini, Paecilini) making them equal by priority rank, but he considers that there is a first revisor, Gill (1893: 133) who gives precedence to Poeciliidae [Gill, T.N. 1893. Families and Subfamilies of Fishes. Memoirs of the National Academy of Science, Mem. VI: 127-138.]

Footnote 12 : Orestiad- as stem derived from feminine (not masculine) gender of genus Orestias root, as proposed in Huber (2005c), confirmed by Cruz-Jofré et al. (2013) [Cruz-Jofré, F., M.A. Valladares, I. Vila & M.A. Mendez. 2013. The genus Orestias (Teleostei: Cyprinodontidae): nomenclatural Errors in the Assignation of species Names. Zootaxa , 3746 (4): 597–599] and reconfirmed by Freyhof et al., 2017 [Freyhof, J., M. Özulug & G. Saç. 2017. Neotype Designation of Aphanius iconii, first Reviser Action to stabilise the Usage of A. fontinalis and A. meridionalis and Comments on the Family Group Names of fishes placed in Cyprinodontidae (Teleostei: Cyprinodontiformes). Zootaxa, 4294 (5): 573-585], without mention of previous works by Huber (2005c) and by Cruz-Jofré et al. (2013); note : in a hitherto overlooked publication [Gill, T. 1896. Note on the Nomenclature of the Poecilioid Fishes. Proc. U.S. Nat. Mus., 18 (1060): 221-224.], Orestiadinae is already proposed, based on its Greek root ; however some researchers still uses Orestiinae or Orestiini as family-group names and Orestiinae or Orestiini are maintained by Laan, Eschmeyer & Fricke (2014), based on subjective prevailing recent practice (however, see foonote 18 on Rachoviini and it is not followed herein) ; on the other hand, Freyhof et al. (2017) [Freyhof, J., M. Özulug & G. Saç. 2017. Neotype Designation of Aphanius iconii, first Reviser Action to stabilise the Usage of A. fontinalis and A. meridionalis and Comments on the Family Group Names of fishes placed in Cyprinodontidae (Teleostei: Cyprinodontiformes). Zootaxa, 4294 (5): 573-585] favor the spelling Orestiini, based on prevailing usage, as they understand it.

Footnote 13 : Anablepsidae and Anablepsinae are the correct names following ICZN requirements derived from genus Anableps root, as formally proposed with argumentation in Huber (2005c) and already listed as such by Fowler (1954) [or equally if another ICZN rule is followed using the entire genus name as a prefix], however this group of fish is mainly viviparous… researchers are different, presently they are still keeping the old incorrect names Anablepidae and Anablepinae, but they may have not been informed of these changes ; Anablepidae and Anablepinae are maintained by Laan, Eschmeyer & Fricke (2014), based on subjective prevailing recent practice (however, see foonote 18 on Rachoviini).

Footnote 14 : Yssolebias has been moved by Costa (2015) to Cyprinodontoids, closer to Cubanichthys, based on the availability of a new, much clearer, radiograph of the single known specimen (holotype of Cyprinodon martae) ; the move has been confirmed by Huber (2015) and the genus has been placed in a new family-group name among Cyprinodontidae as a tribe of Cubanichthyinae to conform with Costa, but with reservations pending further studies.

Footnote 15 : Laan, Eschmeyer & Fricke (2014) have shown that family-group name Rivulidae Myers (1925) based on the genus name Rivulus is preoccupied by Rivulini Grote (1895) that is based on Rivula Guenée in Duponchel (1845), a tribe in Noctuidae (Lepidoptera), and have then proposed the case be referred to the ICZN Commission for a ruling to remove homonymy, following article 55.3.1. (in May 2017 such a request of ruling to ICZN has still not be published, but on August 31. 2017, the following request to ICZN by Laan to ICZN is published [as case 3747: Rivulidae Myers, 1925 (Osteichthyes, Cyprinodontiformes): proposed emendation of the spelling to Rivulusidae to remove homonymy with Rivulinae Grote, 1895 (Lepidoptera, Noctuoidea, Erebidae)], then after review, a new request is published in May 2019 [again as Case 3747 – Rivulinae Grote, 1895 (Lepidoptera, Glossata, Noctuoidea) and Rivulidae Myers, 1925 (Osteichthyes, Cyprinodontiformes): proposed Emendation of the Spelling of the lepidopterous subfamily to Rivulainae to remove Homonymy], this time authored by Laan & Nieukerken, a noted entomologist, proposing the reverse, and notably to maintain Rivulidae for Cyprinodontiformes and in practical terms, the new proposed decision would be definitive if nobody is appealing against the request, say within the next 2 years, but 2 contesting appeals by 2 independent lepidopterists are published late 2019 and ICZN commissioners will have to rule a decision (probably in 2020 or 2021).

Footnote 16 : Costa (2008) creates family-group name Prorivulini with type-genus as Prorivulus and unchanged definitions and diagnoses vs. other family-group name Rivulini, while Huber (2012) in his computerized review of Rivulinae+Kryptolebiatinae places Prorivulus nested in genus Rivulus s.l., closest to subgenera Atlantirivulus and Melanorivulus, implicitly including (synonymizing) Prorivulini into Rivulini, or downgrading it to subtribe level ; pending further evidence in a more comprehensive group of taxa, notably with molecular data on Prorivulus, that synonymization move is not followed herein, however if family-group name Rivulidae would be suppressed by ICZN ruling, then Prorivulini would have a place, at tribe or subtribe level, like Plesiolebi(at)ini, Rachovi(i)ni and Neofundulini ; Costa (2014c) proposes that genus Aphyolebias be synonymized into senior genus Moema, but unfortunately without a new diagnosis of the redefined Moema and without dealing with associated family-group names ; because of lack of re-diagnosis of extended genus Moema and pending further evidence in a more comprehensive group of taxa, that move is partly followed immediately herein (i.e. Aphyolebias downgraded to subgenus of Moema), since such a move is accepted in for publications by authors along description of Aphyolebias new sp. (kenwoodi, juanderibaensis), but on the contrary, family-group name, at subtribe level, Moemina, is herein synonymized into Aphyolebiina -or Aphyolebiatina-, because the former name is created later (i.e. a few paragraphs after, at same page of publication), then is junior, and Aphyolebiina has priority, even if genus Moema has priority over genus Aphyolebias.

Footnote 17 : Huber (2005) proposes family-group names based on correct stems, notably regarding those names built upon genera containing Lebias, with stem as lebi- ; however Laan, Eschmeyer & Fricke (2014) have shown that family-group names should not be corrected based on a more accurate stem for names created after 1999, following article 29.4 and 29.5 (unjustified emendation) : for example correction based on accurate roots of genus name Aphyosemion as Aphyosemiina instead of Aphyosemina is not accepted by them (and provisionally followed herein) ; however corrections based on accurate roots of genus name ending by Lebias, as -lebi (instead of -lebiat) are left open by them for names created before 2000 (except for Kryptolebiatinae because the name has been created after 2000, unlike all other family-group names based on Lebias genus root) and herein the conservative option is followed with -lebiat, like in Cynolebiatinae, up until a formal ICZN-wise publication formally validating Cynolebiinae and others by any author is published (alternatively, the present author herein, as an electronic document, and Laan, one of the above co-authors, who has e-published with ISSN number (2468-9157) his full list of valid fish family-group names starting with «Freshwater Fish List 19th Edition October 2016», both favor Cynolebiinae and similar cases and they should be followed by all authors, even if a formal request and decision by ICZN would help their acceptance) ; debates on the interpretation of those rules, including the date of Killi-Data 2000 (latest rules, for 2000 onwards, or previous ICZN rules because of its closing date is 1999), are still active today and the herein selected options are still temporary and may be reversed in the future) ; notes : on August 31. 2017 the following request to ICZN by Laan is published [as case 3747: proposal for a ruling on the stem of Cynolebias Steindachner, 1876. Richard van der Laan. Bulletin of Zoological Nomenclature] and on April 30. 2019, the following request to ICZN by Laan is published with similar conclusions, opting for stem as -lebi [as case 3793: Laan, R. van der. 2019. Case 3793- Request for a Confirmation of the correct Stem for Family-Group Names proposed before 2000 and typified by fish genera with names ending in -lebias. Bulletin of Zoological Nomenclature, 76: 48-52], and it is herein followed as latest evidence, pending ICZN ruling (yet unpublished in February 2023).

Footnote 18 : Laan, Eschmeyer & Fricke (2014) have used Rachoviina (correctly, based on stem) instead of Rachovina, however Costa started first with Rachovina, then changed it for Rachoviina, then moved back to Rachovina and kept his correction as Rachovini in 2014 [Costa, W.J.E.M. 2014c. Phylogeny and evolutionary Radiation in seasonal rachovine killifishes: biogeographical and taxonomical Implications. Vertebrate Zoology, 64 (2) : 177-192, 1 fig.] ; besides, should Neofundulini and Rachoviini be merged (with a number of valid subtribes, yet undetermined), as mentioned by Laan, Eschmeyer & Fricke (2014), then Neofundulini would have priority over Rachoviini because Costa (1998) have used Neofundulida at a higher rank (infrafamily) over Rachoviidi (supertribe), even if those family-group names (infrafamily, supertribe) are not recognised by ICZN and even if Costa (2014) reverses the situation without argumentation ; in total, if Costa is to be followed, then Rachovini and precedence over Neofundulini should be preferred (and prevailing recent practice) and reversally if Laan et al. are to be followed (as herein, temporarily because of latest evidence and correct stems), then Rachoviini and precedence of Neofundulini should be preferred (and questionable prevailing recent practice, not followed), unlike the case of Anablepsidae, as mentioned in footnote 13.

Footnote 19 : Costa has proposed in 2008 that other family-group names be built from full names of genera, (1) Callopanchacini with a stem based on callopanchac-, (2) Aphyosemionina with a stem based on aphyosemion- ; Laan, Eschmeyer & Fricke (2014) have shown those are un-necessary changes, because original stem is correct (1) or alternatively original stem cannot be changed (resp. 2), even they fit, in part, with ICZN rules.

Footnote 20 : present systematics of Pantanodontidae are herein based on latest evidence by Meinema & Huber, 2023 and by Huber & Meinema, 2024, with morpho-osteology, and by Bragança, Amorim and Costa, 2018 from a multigene molecular analysis of a single component (other congener, not studied), as the most basal unit of suborder Cyprinodontoidei, already initiated by Pohl, Milvertz, Meyer & Vences (2015), who place (today) Aliteranodon ndoano, previously as Pantanodon stuhlmanni in a rogue positioning outside all lampeyes, implying resurrection of family Pantanodontidae (already proposed by Rosen in 1965, upgraded from subfamily level by Whitehead in 1962) ; on the other hand, in the same publication, from molecular data, Bragança, Amorim and Costa spin-off all other lampeyes from Poeciliidae, and upgrade them as 2 distinct families, Procatopodidae for African components and Fluviphylacidae for Southamerican components, but this is based only on 6 of those lampeye species, out of about 70) and that is not followed herein ; the in-depth analysis of that paper raises several questions that push to post-pone those changes, other than Pantanodon, until further confirmations and to let time to time : (1) too few species of lampeyes are included in the study, notably none from East Africa, only one from Central Africa, (2) no species of lampeyes outside the deep-bodied group (Procatopus and related genera, Aplocheilichthys) are included in the study, (3) no species of Cubanichthyinae is included in the study (while following morpho-osteology, they would be positioned in the tree exactly in-between Aliteranodon ndoano (previously as Pantanodon stuhlmanni) and Cyprinodontidae, (4) the authors do not consider alternative (more lumping) strategies, in a more parsimonious process, to define families with new components diagnosed and with subfamilies, ending up with less numerous families (and new diagnoses), (5) philosophically if the authors claim a splitting strategy for their 5 families (moving the previous 7 or 8 families in Cyprinodontiformes to 14), which is respectable, they contradictorily propose a lumping strategy for Aplocheilidae, in one family without explanation, instead of 3, Aplocheilidae, Nothobranchiidae and Rivulidae, well separated by previous molecular data ; at this stage, latest morphological evidence by Ghedotti (2000) (and just before by Huber, 1999) are totally in-congruent with molecular evidence for lampeyes (also, a general situation to all fishes) and more research is needed to clarify that unsatisfactory picture, all the more that Bragança (pers. cmm to Huber, May 2019) announces a new phylogeny of lampeyes based on a much larger sample, in the future ; in total herein, apart from upgrading African lampeyes (that are spun-off from genus Pantanodon) to full family status as Procatopodidae, and separating Congopanchax as a distinct genus based on a later survey, all lower levels of that family are necessarily kept unchanged until missing data and further stable evidence (expected by Bragança in 2022) for all generic components are published, including Aplocheilichthys (which is nested nearby Procatopus, hence with Aplocheilichthyinae as a junior synonym of Procatopodinae).

Footnote 21 : present systematics of Goodeinae are unstable (but strongly studied) with as much as about 2 dozens of valid genera and about 3 dozens of valid extant species, molecularly more or less confirmed, showing strikingly that those fish are strongly differentiated in terms of morphology ; however recent studies have shaded previous results showing that some divided morphospecies are in fact ecotypes of a single phylogenetic species ; besides, some genera are considered as valid by some authors and synonyms for some others (e.g., Hubbsina, Neotoca, the former herein as synonym, because unstable in molecular trees, the latter as valid), and, independently, genus Xenotoca is molecularly paraphyletic but the description of a new genus, if necessary, is pending since at least a decade ; according to Parker et al. (2019), molecular results for all genera and family-group names in Goodeidae based on nuclear data are not consistent with molecular results based on mitochondrial data on both upper nodes and lower nodes (and genus Ataeniobius is weakly supported) ; in total the herein given picture is the mirror of present, but still quite unstable, evidence in terms of the number of valid genera (it could be reduced to a few… but with new uneasy diagnoses to be done) and of the valid family-group names [ref.: Parker, E., A. Dornburg, O.D. Dominguez & K.R. Piller. 2019. Assessing Phylogenetic Information to Reveal Uncertainty in Historical Data: An Example Using Goodeinae (Teleostei: Cyprinodontiformes: Goodeidae). Mol. Phylogenet. Evol., doi.org/10.1016/j.ympev.2019.01.025] ; the synonymization of Chapalichthyini into Zoogoneticini is published in Huber (2019) following molecuar evidence of Chapalichthys nested sub-branch.

Footnote 22 : present systematics of Poeciliidae are stable based both on male gonopodium detailed morphology and on molecular data, except 3 issues : (1) the lumping or not of all previous subgenera of Poecilia into a single genus and at least 9 subgenera, or, the reverse, the splitting of nearly all valid subgenera into distinct genera, the former option being retained herein because it corresponds to consensus of authors, substantiated by latest molecular evidence ; (2) the genus or subgenus status of genera in Heterandriini, notably genera related to Brachyrhaphis, Priapichthys, Heterandria, the latter option as most valid being retained herein because it corresponds to latest molecular evidence, even if that option does not meet consensus between authors ; (3) major differences occurs between morpho-osteological studies using cladistics and molecular data, e.g., Parenti & Rauchenberger, 1989 list under Heterandriini, Heterandria, including Pseudoxiphophorus, Priapichthys, Neoheterandria, Poeciliopsis, Phallichthys, or they list under Poeciliini, Alfaro, Poecilia, Priapella, Xiphophorus, or they list under Gambusiini, Gambusia, Belonesox and Brachyrhaphys, or Lucinda & Reis separate Phalloptychus from rest of Cnesterodontini, but herein in Poeciliidae molecular data are followed instead ; besides family-group name Belonesocini is older than Gambusiinae, then it has priority despite consensus of authors, according to Huber (in print) and Lebistes poecilioides is a junior synonym of Poecilia vivipara, inducing synonymization of Lebistes into Poecilia s.s., but there is no strong consensus on that issue [ref.: Reznick D.N., A.I. Furness, R.W. Meredith & M.S. Springer. 2017. The Origin and biogeographic Diversification of fishes in the family Poeciliidae. PLoS ONE, 12 (3): e0172546. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0172546] .

Footnote 23 : first, but preliminary, analysis based on full mitogenome, discloses that Aphaniidae is probably ambiguous and that its validation still needs more phylogenetic supports ; unfortunately there is still no publication of full mitogenome of Valencia, then for Valenciidae, at this time, in order to clarify at least fusion or clarification of those 2 families [ref.: Teimori, A. & M. Motamedi. 2019. The First Complete Mitochondrial Genome Sequence in the Genus Aphanius (Teleostei). Jounal of Ichthyology, 59 (5): 754–765.] ; and this is congruent with consensus molecular pyramid of upper fish groups (from family to top), regularly updated, by Betancur et al. [latest edition : Phylogenetic Classification of Bony fishes, BMC Evol. Biol., 2017, 17, 162. https://bmcevolbiol.biomedcentral.com/articles/10.1186/s12862-017-0958-3] ; 2 contemporary publications on Aphaniidae systematics are strongly contradictory in their results : Esmaeili et al. (2020), based on new genetic evidence, consider the split of Aphanius in its since long sense to be limited to 3 genera, with redefined genus Aphanius having 5 subgenera or species-groups whereas Freyhof & Yogurtçuoglu (2020) propose the full split of Aphanius into 8 distinct genera, but that is based on no new evidence and rather on a compact opinion of genus concept, then Killi-Data follows the first (former) publication, up to further new evidence (latest development is from Lopeez et al. [Lopez, A.S., T.L. Nester, S. Perea & I. Doadrio. 2023. Complete mitochondrial Genome of the Spanish toothcarp, Aphanius iberus (Valenciennes, 1846) (Actinopterygii, Aphaniidae) and its phylogenetic position within the Cyprinodontiformes order Molecular Biology Reports, https://doi.org/10.1007/s11033-022-08236-w] who favor {like herein} within ICZN conservative ethics to maintain Aphanius as the valid genus in Spain, not Apricaphanius.

Footnote 24 : Amorim & Costa (2022) clearly show with molecular data (and strong bootstrap values) that a component of this subgenus Benirivulus is strong related to the micropus complex (including atratus superspecies), therefore reinforcing the synonymous status of Benirivulus but their work is based on a small group of taxa and if the genus Rivulus s.l. is fully split (much more than presently) then it is unclear if Benirivulus remains still synonymous and to what taxon (Anablepsoides being heterogeneous) or revalidated [ref. : Amorim, P.F. & W.J.E.M. Costa 2022. Evolution and Biogeography of Anablepsoides killifishes shaped by Neotropical geological Events (Cyprinodontiformes, Aplocheilidae). Zoologica Scripta, 53, https://onlinelibrary.wiley.com/doi/10.1111/zsc.12539.

 

 

Important comments on Cyprinodontiformes Tree of Life 

 

Important comments on family-group names and molecular tree of life (American project : https://fishtreeoflife.org/taxonomy/order/Cyprinodontiformes/) : the above herein (artificial) list of family-group names is the mirror of latest published evidence in Killi-Data (the tree of life project is much delayed with latest update as 2017, due to slow checking and homogeneization processes on all groups of living beings on Earth) ; at family-group names levels, currently, molecular data have taken precedence over cumulative knowledge with (historically) morphological, micromorphological, osteological, cladistic and computer combined techniques ; 3 waves of molecular knowledge have occurred : (1) early days (1993… ) with limited sequences and limited sampling (aiming at understanding fish groups evolutions and dated origin with plates tectonics and major palaeo-climate events, (2) maturity period (ca. 2000-2015… ) with multiple but still limited sequences and a lot of sampling, still uncomplete (culminating for Cyprinodontiformes, with all groups being evaluated, in 2020, last with lampeyes), (3) the extension period using full genome (nuclear and mitochondrial) but with slowly iincreasing number of samples (due to still high costs) aiming to mirror true tree of life, mainly with fossils (few at the beginning, then more numerous, knowing that fossils are very useful but have the set-back to indicate a minimum age, not the precise correct age) and palaeo-plate tectonics as comparative calibrations ; recent major molecular studies (2017) on probable ages and inter-relationships of higher groups of fish suggest that increased gene sampling, in general, might have a greater beneficial effect on the rigor of the estimation of phylogenetic topologies than more extensive taxon sampling (in short, large DNA fragments give better results, hence the few used taxa, i.e. those fish used as general fish models like Salmon, Carp, and Danio rerio, Fundulus heteroclitus, Oryzias latipes, Tetraodon sp. with Homo sapiens as the outgroup) ; in the resulting (preliminary) tree, old-old ancestor of Killifish have separated from pufferfish about 190 Million years ago (= MYA) (at about the time when Pangaea split, but with very high uncertainties in terms of times brackets), and are related to Beloniformes (themselves separated a "little" later), and to Perciformes (cichlids, the "younger" group, separated about 113 MYA, therefore after the split of Gondwana and the drift of present Africa and South America)… but are not related to Cyprinids (Barbs), to Siluriformes or to Salmon (anecdotally Cypriniformes, i.e. Barbs, and Cyprinodontiformes, i.e. Killifishes, were thought closely related by early zoologists about 150 years ago, only separated by the presence of teeth in the latter !) ; further, the monophyly (with a single ancestor) of superorder Atherinomorphae (Atheriniformes, Beloniformes, and Cyprinodontiformes, i.e. medakas, ricefishes, flyingfishes, silversides, rainbowfishes, killifishes) is confirmed on whole mitogenome sequences (not nuclear) with their relationship to Cichlomorphae (including Cichliformes or cichlids and other groups that are in part spawning demersal eggs with filaments), in agreement with the widely accepted morpho-osteological hypothesis that ricefishes (family Adrianichthyidae) are more closely related to Belonoids than to Cyprinodontiformes [Refs. : Steinke, Dirk ; Salzburger, Walter ; Meyer, Axel. Journal of Molecular Evolution, 2006 June; 62 (6): 772-784, 4 figs.|Setiamarga, Davin H.E., Miya Masaki, Yamanoue Yusuke, Mabuchi Kohji, Satoh Takashi P., Inoue Jun G., Nishida Mutsumi, Molecular Phylogenetics and Evolution, 49 (2008) 598–605] ; on Cyprinodontiformes alone, there is also new evidence (and dictinct, less old, palaeo-timings, based on multiple fossils) with paper by Reznick et al. 2017 [ref. Reznick D.N., A.I. Furness, R.W. Meredith & M.S. Springer. 2017. The Origin and biogeographic Diversification of fishes in the family Poeciliidae. PLoS ONE, 12 (3): e0172546. doi:10.1371/journal. pone.0172546], showing probable age of Cyprinodontiformes as 90.7 MYA using independent rates ; very recently (2020), using full genome and better dated fossils, molecular research dates first origin of both Cichlidae and Cyprinodontiformes not very distantly to about 70 MYA, rather similar (with obvious uncertainty) to 66 MYA, the K/T boundary, when a giant meteor slammed into Earth off the coast of modern-day Mexico, then killed nearly everything on Earth during an apocalyptic turmoil lasting ca. 5 MYA, but the (Swiss-based) authors still discuss the 3 previous scenarios to explain vicariance-dispersal schemes for fishes and notably the 2 mainly tropical groups, Cichlidae and Cyprinodontiformes, because both are secondary freshwater fishes (i.e., presently mainly living in freshwaters, but historically living in marine (coastal ?) waters [ref.: Matschiner, M., Böhne, A., Ronco, F., Salzburger W. The genomic timeline of cichlid fish diversification across continents. Nature Communications, 11, 5895 (November 2020), freely readable online at https://www.nature.com/articles/s41467-020-17827-9] ; even more recently, a mainly Japanese large team studying Adrianichthyidae (a family holding Oryzias and Adrianichthys in Beloniformes) with full mitogenomes and parts of nuclear genomes and fossil calibration, show that Cyprinodontiformes origin may lie at ca. 80-85 MYA with a blue confidence interval that extends +/- 15 MYA, and first direct ancestor emergence at ca. 90-95 MYA, more in line with Reznick et al. (2017) [Ref. : Kazunori et al., 2021 : Mesozoic Origin and ‘out-of-India’ radiation of ricefishes (Adrianichthyidae). Biology Letters 17: 7 pp., suppl., https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0212]. Then, to sum-up, the most recent evidence for dating origin of Cyprinodontiformes points out 4 alternate scenarios : (1) vicariance mainly when Gondwana is a single unsplit continent (ca. 150 to 120 MYA), a scenario pushed by early molecularists that is nicely congruent with continent drifts, but not at all with fossil dating, (2) subsequent pseudo-vicariance, ca. 100-90 MYA, when Gondwana is weakly split with Atlantic ocean limited to a 'narrow and shallow' (very relatively) channel between South America and Africa (and similarly between East Africa and India-Madagascar) and when Atherinomorphae and Cyprinodontiformes as coastal marine fish can easily extend, then migrate into nearby brackish, then fresh waters (in line with Reznick et al., Kazunori et al.), (3) oceanic dispersal contemporary (to K-T boundary, 65 MYA), in fact a 'little' (by MYA !) anteriorly or posteriorly to the meteor catastrophy in line with an option by Matschiner et al. but not in accordance with known fossils of Cyprinodontiformes and not in accordance with the missing capacity (as far as known) for Cyprinodontiformes of migrating thru deep seas over thousands of kms, or (4) much later independent multiple colonizations (i.e. ca 45 MYA) more in line with another option by Matschiner et al., some partial molecular data and fossil evidence of Cyprinodontiformes but not in accordance with the availability of only 1 or 2 ancestor(s) per continent, for India, Madagascar, Africa, South America (why not more ?), in line with dispersal along floating seaweed mats or tiny land rafts at sea surface (a theory publicized by scientist Alan de Quieroz, supported once by Brasilian Wilson Costa), putatively in correspondence with present major family-group names in the tropics, but not in accordance with previous key role of palaeo-Tethys sea of first scenario… latest molecular research with known fossils would favor scenario n°2 for the emergence of ancestor of Cyprinodontiformes (85-100 MYA), when Atherinomorphae expanded all over coasts of just fully split Gondwana, before, unlike most its other components, Cyprinodontiformes decide to go for freshwaters (when exactly ? along a single move or more probably several ? from coastal fringes or from deep seas ?)… future will tell more, hopefully, on that fascinating enigma of origin, type of move-expansion and initial timing of emergence-breakups of Cyprinodontiformes families (see below sketches to illustrate the 4 scenarios). Beware, those 4 scenarios are theoretical and speculative for Cyprinodontiformes and like for most fish groups, emergence (order level), first breakup (family level) and further branches of tree (subfamily level) are presently very unstable-uncertain (for more on palaeo-tectonics look at ODSN over time (e.g., type Age to be reconstructed [My]: 110, or 100, or 95, or 90, or 85, up to K-T boundary 65, or 45, and so on). Note : research is going on aiming at a an appropriate date of emergence of Atherinomorphae and Cyprinodontiformes, the latest, based on 12 molecular markers, by Morales et al. (2024), showing that the separation of the former from upper branches is dated before the K-T boundary (65 MYA), i.e. a bit before in terms of MYA, and the separation of the latter is dated after the K-T boundary (65 MYA), i.e. a bit after in terms of MYA (cf. https://doi.org/10.1186/s12864-024-10416-w).

 

 

 

 

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