Dorsum of Leptotocepheus trimucronatus, from Balogh & Balogh (1992 vol. 2).

Belongs within: Brachypylina.
Contains: Nippobodidae, Dampfiella, Beckiella, Odontocepheus (Odontocepheus), Diplobodes, Yoshiobodes, Gibbicepheus (Gibbicepheus), Carabodinae, Fissicepheus (Fissicepheus), Pseudotocepheus (Pseudotocepheus), Plenotocepheus, Otocepheinae, Dolicheremaeus.

The Carabodoidea are a primarily tropical group of oribatid mites, many of which spend as least part of their life as burrowers in decaying wood. Members of this group include the Carabodidae, characterised by more or less parallel lamellae on the prodorsum (Balogh & Balogh 1992). Though diverse in adult biology, all species are probably endophagous as nymphs. Adults of many species are arboreal, feeding on fungi and lichens (Norton & Behan-Pelletier 2009). In a number of genera the notogaster bears longitudinal rows of large tubercles, such as in Gibbicepheus or Diplobodes. The genera Spathulocepheus and Trichocarabodes are distinctive in the high number of setae on their genital plates: eight pairs in Trichocarabodes, ten in Spathulocepheus. Other genera have at most seven pairs of genital setae (Balogh & Balogh 1992).

Members of the family Dampfiellidae have the tibiae and tarsi fused and moving as a single segment. The Otocepheidae are characterised by well-developed pedotecta (outgrowths of the body wall underlying the base of the legs). Members of this family are particularly common in damp habitats such as moist litter (Norton & Behan-Pelletier 2009). The family is sometimes divided between two families, Otocepheidae and Tetracondylidae, on the basis of the development of the pedotecta. In Otocepheidae sensu stricto, pedotecta 2+3 are particularly large, so that they are axe-shaped in ventral view or tailfin-shpaed in dorsal view. In 'Tetracondylidae', the pedotecta are distinct but not so extremely developed (Balogh & Balogh 1992).

Carabodoidea [Otocepheoidea] S04
    |  i. s.: Nippobodidae S04
    |         Tokunocepheus Aoki 1966 [Tokunocepheidae] S04
    |           `--*T. mizusawai Aoki 1966 S04
    |         Carabocepheus Berlese 1910 [Carabocepheidae] S04
    |           `--*C. lounsburyi (Berlese 1910) [=Carabodes (*Carabocepheus) lounsburyi] S04
    |                |--C. l. lounsburyi S04
    |                `--C. l. latior Balogh & Mahunka 1966 S04
    |--Dampfiellidae S04
    |    |--Dampfiella S04
    |    `--Beckiella PL17
    `--+--Carabodidae SK10
       |    |  i. s.: Apotomocepheus Aoki 1965 S04
       |    |           `--*A. gressitti Aoki 1965 S04
       |    |         Meriocepheus Aoki 1973 S04
       |    |           `--*M. peregrinus Aoki 1973 S04
       |    |         Sagittabodes Balogh & Balogh 1992 S04
       |    |           `--*S. sagitta (Balogh & Mahunka 1966) [=Machadocepheus sagitta] S04
       |    |         Tansocepheus Mahunka 1983 S04
       |    |           `--*T. serratus Mahunka 1983 S04
       |    |         Tectocarabodes Mahunka 1988 S04
       |    |           `--*T. monstruosus Mahunka 1988 S04
       |    |         Tuberocepheus Balogh & Mahunka 1969 S04
       |    |           `--*T. longus (Balogh 1964) [=Machadocepheus longus] S04
       |    |         Singabodes Mahunka 1998 S04
       |    |           `--*S. rarus Mahunka 1998 S04
       |    |         Archegocepheus Aoki 1965 S04
       |    |           |--*A. imadatei Aoki 1965 S04
       |    |           `--A. macrofoliatus Wen 1997 S04
       |    |         Bathocepheus Aoki 1978 [incl. Philippobodes Balogh & Balogh 1992] S04
       |    |           |--*B. concavus Aoki 1978 S04
       |    |           `--B. manguiati (Corpuz-Raros 1979) S04 (see below for synonymy)
       |    |         Cavernocarabodes Mahunka 1974 S04
       |    |           |--*C. perreti Mahunka 1974 S04
       |    |           `--C. trigonosternum (Pérez-Íñigo 1976) [=Carabodes trigonosternum] S04
       |    |         Spathulocepheus Balogh & Mahunka 1969 S04
       |    |           |--*S. amazonicus Balogh & Mahunka 1969 S04
       |    |           `--S. plumiger Balogh & Mahunka 1977 S04
       |    |         Malgasodes Mahunka 2000 S04
       |    |           |--*M. curvisetus Mahunka 2000 S04
       |    |           `--M. hungarorum Mahunka 2000 S04
       |    |         Gymnobodes Balogh 1965 S04
       |    |           |--*G. fraterculus (Balogh 1963) [=Carabodes fraterculus] S04
       |    |           |--G. semengok Mahunka 1996 S04
       |    |           `--G. subnudus (Balogh 1963) [=Carabodes subnudus] S04
       |    |         Pseudocarabodes Mahunka 1991 [incl. Bakobodes Mahunka 1996, Yemenobodes Mahunka 2000] S04
       |    |           |--*P. xenus Mahunka 1991 S04
       |    |           |--P. improvisus (Mahunka 2000) [=Yemenobodes improvisus] S04
       |    |           `--P. orangutan (Mahunka 1996) [=Bakobodes orangutan] S04
       |    |         Trichocarabodes Balogh 1961 [incl. Ceylobodes Balogh & Balogh 1992] S04
       |    |           |--*T. celisi (Balogh 1958) [=Carabodes celisi] S04
       |    |           |--T. capillatus Balogh 1970 S04 [=*Ceylobodes capillatus BB92]
       |    |           |--T. costulatus Mahunka 1984 S04
       |    |           `--T. hettigei Balogh 1970 S04
       |    |         Hardybodes Balogh 1970 [incl. Carabodella Mahunka 1986] S04
       |    |           |--*H. mirabilis Balogh 1970 S04
       |    |           |--H. calcaratus (Mahunka 1986) S04 [=*Carabodella calcarata BB92]
       |    |           |--H. flabellatus Mahunka 1995 S04
       |    |           `--H. penicillatus Mahunka 1995 S04
       |    |         Cubabodes Balogh & Mahunka 1974 S04
       |    |           |--*C. hexagonalis Balogh & Mahunka 1974 S04
       |    |           |--C. confertus Balogh & Mahunka 1980 S04
       |    |           |--C. radiatus Balogh & Mahunka 1974 S04
       |    |           |--C. spathulatus Balogh & Mahunka 1980 S04
       |    |           `--C. verrucatus Balogh & Mahunka 1980 S04
       |    |         Machadocepheus Balogh 1958 S04
       |    |           |--*M. excavatus Balogh 1958 S04
       |    |           |--M. filiferus Mahunka 1986 S04
       |    |           |--M. pauliani Balogh 1962 S04
       |    |           |--M. phyllophorus Balogh 1970 S04
       |    |           |--M. taprobanicus Balogh 1970 S04
       |    |           `--M. tuberculosus Mahunka 1987 S04
       |    |         Odontocepheus Berlese 1913 S04
       |    |           |--O. (Odontocepheus) S04
       |    |           |--O. (Aokiella Balogh & Mahunka 1967) S04
       |    |           |    |--O. (*A.) florens (Balogh & Mahunka 1967) [=*Aokiella florens] S04
       |    |           |    `--O. (A.) rotundus (Hammer 1979) [=Aokiella rotunda] S04
       |    |           `--O. (Indotocepheus Mondal & Kundu 1999) S04
       |    |                `--O. (*I.) himalayensis (Mondal & Kundu 1999) [=*Indotocepheus himalayensis] S04
       |    |         Pasocepheus Aoki 1976 S04
       |    |           |--P. (Pasocepheus) [incl. Opisthocepheus Aoki 1976] S04
       |    |           |    |--*P. (P.) triarcuatus Aoki 1976 S04
       |    |           |    |--P. (P.) bako Mahunka 1996 S04
       |    |           |    |--P. (P.) eremaeozetoides Mahunka 1995 S04
       |    |           |    `--P. (P.) kirai (Aoki 1976) S04 [=*Opisthocepheus kirai BB92]
       |    |           `--P. (Guineobodes Mahunka 1987) S04
       |    |                `--P. (*G.) papuanus (Balogh 1970) [=Machadocepheus papuanus] S04
       |    |         Congocepheus Balogh 1958 S04
       |    |           |--*C. heterotrichus Balogh 1958 S04
       |    |           |--C. hauseri Mahunka 1989 S04
       |    |           |--C. involutus Mahunka 1997 S04
       |    |           |--C. latilamellatus Mahunka 1984 S04
       |    |           |--C. orientalis Mahunka 1989 S04
       |    |           |--C. ornatus Mahunka 1983 S04
       |    |           |--C. taurus Balogh 1961 S04
       |    |           `--C. velatus Mahunka 1986 S04
       |    |         Diplobodes S04
       |    |         Yoshiobodes S04
       |    |--Gibbicepheus Aoki 1958 S04 [Gibbicepheinae CH98]
       |    |    |--G. (Gibbicepheus) S04
       |    |    `--G. (Gibbibodes Mahunka 1986) S04
       |    |         `--G. (*G.) similis (Mahunka 1986) [=*Gibbibodes similis] S04
       |    `--Carabodinae PL17
       `--Otocepheidae [Tetracondylidae] S04
            |  i. s.: Ocellotocepheus Mahunka 1989 S04
            |           `--*O. tuberculatus Mahunka 1989 S04
            |         Samarocepheus Corpuz-Raros 1990 S04
            |           `--*S. cavernicolus Corpuz-Raros 1990 S04
            |         Cerostocepheus Mahunka 1973 S04
            |           `--*C. trisetosus (Balogh 1970) [=Eurostocepheus trisetosus] S04
            |         Ikarotocepheus Mahunka 1988 S04
            |           `--*I. alatus Mahunka 1988 S04
            |         Philippotocepheus Balogh & Balogh 1992 S04
            |           `--*P. siniloanensis (Corpuz-Raros 1979) [=Papuacepheus siniloanensis] S04
            |         Borneremaeus Mahunka 1991 S04
            |           `--*B. hauseri Mahunka 1991 S04
            |         Bulbocepheus Mahunka 1988 S04
            |           `--*B. hauseorum Mahunka 1988 S04
            |         Fernandocepheus Mahunka 1982 S04
            |           `--*F. franzi Mahunka 1982 S04
            |         Flagellocepheus Balogh 1984 S04
            |           `--*F. sagittatus Balogh 1984 S04
            |         Hydroecocepheus Corpuz-Raros 1979 S04
            |           `--*H. romeoi Corpuz-Raros 1979 C-R79
            |         Lophotocepheus Balogh & Balogh 1983 S04
            |           `--*L. simplex Balogh & Balogh 1983 S04
            |         Neotrichocepheus Hammer 1973 S04
            |           `--*N. tongaensis (Hammer 1973) [=Dolicheremaeus (*Neotrichocepheus) tongaensis] S04
            |         Paradolicheremaeus Tseng 1982 S04
            |           `--*P. tabulatus Tseng 1982 S04
            |         Seboetocepheus Mahunka 1985 S04
            |           `--*S. coronatus Mahunka 1985 S04
            |         Trichocondyla Balogh & Balogh 1986 S04
            |           `--*T. multisetosa Balogh & Balogh 1986 S04
            |         Papuacepheus Balogh 1968 S04
            |           |--*P. geminatus (Balogh 1968) [=Megalotocepheus (*Papuacepheus) geminatus] S04
            |           `--P. vagorum Corpuz-Raros 1991 S04
            |         Trichocepheus Balogh & Mahunka 1966 S04
            |           |--*T. lamellatus Balogh & Mahunka 1966 S04
            |           `--T. auriculus Grobler 1995 S04
            |         Wallworkodes Balogh & Balogh 2002 S04
            |           |--*W. carinatus (Wallwork 1977) [=Carabodes carinatus] S04
            |           `--W. hyalinus (Wallwork 1977) [=Carabodes hyalinus] S04
            |         Kalayaan Corpuz-Raros 1998 S04
            |           |--*K. bonifacioi Corpuz-Raros 1998 S04
            |           |--K. centenarius Corpuz-Raros 1998 S04
            |           `--K. rizali Corpuz-Raros 1998 S04
            |         Rimandocepheus Corpuz-Raros 1998 S04
            |           |--*R. leoi Corpuz-Raros 1998 S04
            |           |--R. brevisetosus Corpuz-Raros 1998 S04
            |           `--R. longisetosus Corpuz-Raros 1998 S04
            |         Afrotocepheus Mahunka 1985 S04
            |           |--*A. (Afrotocepheus) sinarmatus Mahunka 1985 S04
            |           `--A. (Didierotocepheus Mahunka 1994) S04
            |                `--A. (*D.) berndi (Mahunka 1994) [=*Didierotocepheus berndi] S04
            |         Trichotocepheus Aoki 1965 S04
            |           |--*T. erabuensis Aoki 1965 S04
            |           |    |--T. e. erabuensis S04
            |           |    `--T. e. modestus Aoki 1973 S04
            |           `--T. amamiensis Aoki 1965 S04
            |         Fissicepheus Balogh & Mahunka 1967 S04
            |           |--F. (Fissicepheus) S04
            |           `--F. (Psammocepheus Aoki 1970) S04
            |                |--F. (*P.) amabilis Aoki 1970 S04
            |                `--F. (P.) haradai Choi 1986 S04
            |         Longocepheus Balogh & Mahunka 1966 S04
            |           |--*L. australis Balogh & Mahunka 1966 S04
            |           |--L. globosus Grobler 1995 S04
            |           |--L. longus (Balogh 1961) [=Pseudotocepheus longus] S04
            |           `--L. youngai (Mahunka 1984) [=Nesotocepheus youngai] S04
            |         Spinotocepheus Hammer 1981 S04
            |           |--*S. javensis Hammer 1981 S04
            |           |--S. foveolatus Hammer 1981 S04
            |           |--S. nigromaculatus Hammer 1981 S04
            |           `--S. tjibodensis Hammer 1981 S04
            |         Cavernocepheus Balogh & Mahunka 1969 S04
            |           |--*C. monstruosus Balogh & Mahunka 1969 S04
            |           |--C. acutus Balogh & Palacios-Vargas 1997 S04
            |           |--C. furcatus Balogh & Palacios-Vargas 1997 S04
            |           |--C. fusifer Balogh & Palacios-Vargas 1997 S04
            |           |--C. monticola Balogh 1989 S04
            |           |--C. obliquus Balogh 2002 S04
            |           `--C. undulatus Balogh 2002 S04
            |         Papillocepheus Balogh & Mahunka 1966 [incl. Clavazetes Hammer 1966] S04
            |           |--*P. heterotrichus Balogh & Mahunka 1966 S04
            |           |--P. areolatus Mahunka 1987 S04
            |           |--P. decoratus Mahunka 1994 S04
            |           |--P. decorus (Hammer 1966) S04 [=*Clavazetes decorus BB92]
            |           |--P. deficiens Balogh & Balogh 1983 S04
            |           |--P. neotropicus (Balogh 1988) [=Clavazetes neotropicus] S04
            |           |--P. reductus Mahunka 1994 S04
            |           `--P. tuberculatus (Mahunka 1978) [=Clavazetes tuberculatus] S04
            |         Pseudotocepheus Balogh 1961 S04
            |           |--P. (Pseudotocepheus) S04
            |           `--P. (Constrictocepheus Grobler 1998) S04
            |                |--P. (*C.) geminatus Balogh & Mahunka 1969 S04
            |                |--P. (C.) coarctatus Balogh 1985 S04
            |                |--P. (C.) curtiseta Hammer 1966 S04
            |                |--P. (C.) gobletus Chakrabarti & Mondal 1978 S04
            |                |--P. (C.) monteithi Balogh & Balogh 1983 S04
            |                |--P. (C.) orientalis Mondal & Kundu 1983 S04
            |                |--P. (C.) punctatus Hammer 1966 S04
            |                `--P. (C.) tenuiseta Hammer 1966 S04
            |         Plenotocepheus PL17
            |--Otocepheinae C-R79
            `--Tetracondylinae C-R79
                 |--Dolicheremaeus NB-P09
                 `--Leptotocepheus Balogh 1961 C-R79, S04
                      |--*L. trimucronatus Balogh 1961 S04
                      |--L. flagellifer Mahunka 1988 S04
                      |--L. macromucronatus Mahunka 1988 S04
                      `--L. orientalis Mahunka 1988 S04

Bathocepheus manguiati (Corpuz-Raros 1979) S04 [=Machadocepheus manguiati S04, *Philippobodes manguiati BB92]

*Type species of generic name indicated


[BB92] Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[CH98] Colloff, M. J., & R. B. Halliday. 1998. Oribatid Mites: A catalogue of Australian genera and species. CSIRO Publishing: Collingwood.

[C-R79] Corpuz-Raros, L. A. 1979. Philippine Oribatei (Acarina). I. Preliminary list of species and descriptions of forty new species. Philippine Agriculturist 62 (1): 1–82.

[NB-P09] Norton, R. A., & V. M. Behan-Pelletier. 2009. Suborder Oribatida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 430–564. Texas Tech University Press.

[PL17] Pachl, P., A. C. Lindl, A. Krause, S. Scheu, I. Schaefer & M. Maraun. 2017. The tropics as an ancient cradle of oribatid mite diversity. Acarologia 57 (2): 309–322.

[SK10] Schäffer, S., S. Koblmüller, T. Pfingstl, C. Sturmbauer & G. Krisper. 2010. Ancestral state reconstruction reveals multiple independent evolution of diagnostic morphological characters in the "Higher Oribatida" (Acari), conflicting with current classification. BMC Evolutionary Biology 10: 246.

[S04] Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.


Dorsal (left) and lateral views of Beklemishevia barbata, from Balogh & Balogh (1992).

Belongs within: Palaeosomata.

The Ctenacaroidea are a group of poorly sclerotised oribatids with tridactylous leg tarsi and usually more or less expanded bothridial setae (Norton & Behan-Pelletier 2009).

Characters (from Norton & Behan-Pelletier 2009): Naso absent. Bothridial seta slightly to greatly expanded (rarely filfirm). Larger exobothridial seta (xa) positioned anterior to other (xp), usually about half length of seta in or less. Hysterosoma moderately to much elongated, with aesthenic zone. Opisthonotum with weakly definable notogaster; posterior shield bearing all setae posterior to row c, with latter on separate anterior sclerite; separation between sclerites narrow or broad. Opisthonotum usually neotrichous in pygidial region; rarely with 16 pairs (p4 absent). Setae of various form, large and darkly pigmented or not, d2 and e1 erectile or not; setae h1 and p1 not unusually short. Cupules not evident. Anterior pair of genital papillae greatly reduced or absent; most anterior pair of genital setae normal or strongly modified. Famulus of tarsus I much smaller than solenidia; inserted at midpoint or in distal half of tarsus. Tibia II with one solenidion. Leg pretarsi II–IV tridactylous, with empodial claw well developed or much reduced. Adoral seta or1 strongly barbed or pectinate, usually larger than other two setae. Chelicerae chelate-dentate, oriented obliquely, anteroventrally; digits not narrowed. Palptarsus elongated, at least twice tibial length; eupathidia smooth.

Ctenacaroidea NB-P09
    |--Adelphacarus Grandjean 1952 S04 [Adelphacaridae NB-P09]
    |    `--*A. sellnicki Grandjean 1952 S04
    |--Aphelacarus Grandjean 1932 S04 [Aphelacaridae NB-P09, Aphelacarinae]
    |    `--*A. acarinus (Berlese 1910) [=Parhypochthonius acarinus] S04
    |         |--A. a. acarinus [incl. A. rossicus Zachvatkin 1945] S04
    |         `--A. a. sahariensis Gil-Martín, Subías & Arillo 1992 S04
    `--Ctenacaridae NB-P09
         |--Gilarovella Lange 1974 S04
         |    `--*G. demetrii Lange 1974 S04
         |--Neoctenacarus Moritz 1974 S04
         |    `--*N. hastilis Moritz 1974 S04
         |--Ctenacarus Grandjean 1939 [=Grandjeanacarus Zachvatkin 1945] S04
         |    |--*C. araneola (Grandjean 1932) S04 (see below for synonymy)
         |    `--C. foliisetosus Bulanova-Zachvatkina 1980 S04
         `--Beklemishevia Zachvatkin 1945 S04
              |--*B. galeodula Zachvatkin 1945 S04
              |--B. africana (Mahunka 1974) [=Ctencarus africanus] S04
              |--B. barbata (Schubart 1968) [=Ctenacarus barbatus] S04
              |--B. demeteri Mahunka 1984 S04
              |--B. hispaniola Pérez-Íñigo 1997 S04
              `--B. zachvatkini Bulanova-Zachvatkina 1980 S04

*Ctenacarus araneola (Grandjean 1932) S04 [=Palaeacarus araneola S04, C. araneolus CH98, Grandjeanacarus araneola CH98]

*Type species of generic name indicated


Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 2. Plates. Hungarian Natural History Museum: Budapest.

[CH98] Colloff, M. J., & R. B. Halliday. 1998. Oribatid Mites: A catalogue of Australian genera and species. CSIRO Publishing: Collingwood.

[NB-P09] Norton, R. A., & V. M. Behan-Pelletier. 2009. Suborder Oribatida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 430–564. Texas Tech University Press.

[S04] Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.


Nosema apis, a pathogen of bees, from here.

Belongs within: Fungi.

The Microsporidia are a group of anaerobic intracellular parasites, usually found in animals, which penetrate the cells of their hosts by means of an extrusive polar tube.

Characters (from Adl et al. 2012): Obligate intracellular parasites, usually of animals; mitochondria highly reduced to mitosomes; spores with inner chitin wall and outer proteinaceous wall; without kinetosomes, centrioles or cilia; centrosomal plaque; extrusive specialized polar tube for host penetration; reproduction sexual, asexual or both.

<==Microsporidia [Microspora, Microsporida]
    |--Manubrispora C-S99
    |    |--Metchnikovella [Metchnikovellea] AS12
    |    `--Amphiamblys C-S99
    |--Minisporea C-S98
    |    |  i. s.: Buxtehudia C-S98
    |    |         Hessea C-S98
    |    `--Chytridiopsidae BJ02
    |         |--Chytridiopsis C-S98
    |         |--Cryptosporina BJ02
    |         |--Gurleya BJ02
    |         |--Napamichum BJ02
    |         |--Thelohania californica BJ02, BS94
    |         `--Intexta BJ02
    `--Microsporea [Disporea, Pleistophorea] C-S98
         |  i. s.: Spraguea lophii C-S98, C02
         |         Caudospora C-S98
         |         Amblyospora C-S98
         |         Glugea C-S98
         |           |--G. americanus C02
         |           |--G. danilewskyi D56
         |           |--G. heraldi K-M02b
         |           |--G. hertwigi K-M02a
         |           |--G. plecoglossi HH99
         |           `--G. stephani MS98
         |         Cystosporogenes operophterae BJ02
         |--Pleistophora [Pleistophoridae] BJ02
         |    |--P. culicis OH02
         |    |--P. macrozoarcidis K-MC02
         |    `--P. schubergi OH02
         |--Endoreticulatus schubergi BJ02
         `--+--Vittaforma corneae BJ02 [=Nosema corneum N00]
            `--+--+--Enterocytozoon bieneusi BJ02
               |  `--Nucleospora salmonis BJ02
               `--+--Encephalitozoon BJ02
                  |    |--E. cuniculi BJ02
                  |    `--+--E. hellem BJ02
                  |       `--E. intestinalis BJ02 [=Septata intestinalis N00]
                  `--Nosematida OV95
                       |--Telomyxidae OV95
                       |    |--Telomyxa OV95
                       |    `--Issia OV95
                       |         |--I. globulifera OV95
                       |         |--I. singulati Ovčarenko & Vima 1995 OV95
                       |         `--I. trichopterae [=Perezia trichopterae] OV95
                       `--Nosema BJ02
                            |  i. s.: N. algerae L91
                            |         N. connori N00
                            |         N. echinostomi D56
                            |         N. fumiferanae OH02
                            |         N. legeri D56
                            |         N. locustae SM03
                            |--+--+--N. bombycis BJ02
                            |  |  `--N. trichoplusiae BJ02
                            |  `--+--N. furnacalis BJ02
                            |     `--N. granulosis BJ02
                            `--+--N. apis BJ02
                               `--+--+--Vairimorpha necatrix BJ02
                                  |  `--+--N. oulemae BJ02
                                  |     `--+--N. ceranae BJ02
                                  |        `--+--N. vespula BJ02
                                  |           `--+--N. portugal BJ02
                                  |              `--‘Vairimorpha’ lymantriae BJ02
                                  `--Unikaryonidae BJ02
                                       |--Orthosomella BJ02
                                       |--*Unikaryon piriformis Canning, Lai & Lie 1974 BJ02
                                       |--Larssoniella resinellae BJ02
                                       |--Canningia Weiser et al. 1995 non Cookson & Eisenack 1960 (ICBN) BJ02
                                       `--Oligosporidium BJ02
                                            |--O. arachnicolum BJ02
                                            `--O. occidentalis Becnel, Jeyaprakash et al. 2002 BJ02

Microsporidia incertae sedis:
  Microsporidium CM07
    |--M. ceylonensis N00
    |--M. phytoseiuli BJ02
    `--M. weiseri CM07
  Trachipleistophora GLB04
    |--T. anthropophthera N00
    `--T. hominis GLB04
  Antonospora locustae JK06
  Ameson michaelis C-SC96
  Ichthyosporidium hoferi C-SC96, M02
  Brachiola vesicularum N00
  Desportesia AS12
  Edhazardia aedis BJ02
  Perezia pyraustae OH02
  Pseudopleistophorinae OV95

*Type species of generic name indicated


[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[BJ02] Becnel, J. J., A. Jeyaprakash, M. A. Hoy & A. Shapiro. 2002. Morphological and molecular characterization of a new microsporidian species from the predatory mite Metaseiulus occidentalis (Nesbitt) (Acari, Phytoseiidae). Journal of Invertebrate Pathology 79: 163–172.

[BS94] Buss, L. W., & A. Seilacher. 1994. The phylum Vendobionta: a sister group of the Eumetazoa? Paleobiology 20 (1): 1–4.

[C02] Caruso, J. H. 2002. Goosefishes or monkfishes. Family Lophiidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 264–270. Smithsonian Institute Press: Washington.

[C-S98] Cavalier-Smith, T. 1998. A revised six-kingdom system of life. Biological Reviews 73: 203–266.

[C-S99] Cavalier-Smith, T. 1999. Zooflagellate phylogeny and the systematics of Protozoa. Biological Bulletin 196: 393–396.

[C-SC96] Cavalier-Smith, T., & E. E. Chao. 1996. Molecular phylogeny of the free-living archezoan Trepomonas agilis and the nature of the first eukaryote. Journal of Molecular Evolution 43: 551–562.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[GLB04] Gray, M. W., B. F. Lang & G. Burger. 2004. Mitochondria of protists. Annual Review of Genetics 38: 477–524.

[HH99] Hasegawa, M., & T. Hashimoto. 1999. Phylogenetic position of amitochondriate protists in the evolution of eukaryotes. Biological Bulletin 196: 389–392.

[JK06] James, T. Y., F. Kauff, C. L. Schoch, P. B. Matheny, V. Hofstetter, C. J. Cox, G. Celio, C. Gueidan, E. Fraker, J. Miadlikowska, H. T. Lumbsch, A. Rauhut, V. Reeb, A. E. Arnold, A. Amtoft, J. E. Stajich, K. Hosaka, G.-H. Sung, D. Johnson, B. O'Rourke, M. Crockett, M. Binder, J. M. Curtis, J. C. Slot, Z. Wang, A. W. Wilson, A. Schüßler, J. E. Longcore, K. O'Donnell, S. Mozley-Standridge, D. Porter, P. M. Letcher, M. J. Powell, J. W. Taylor, M. M. White, G. W. Griffith, D. R. Davies, R. A. Humber, J. B. Morton, J. Sugiyama, A. Y. Rossman, J. D. Rogers, D. H. Pfister, D. Hewitt, K. Hansen, S. Hambleton, R. A. Shoemaker, J. Kohlmeyer, B. Volkmann-Kohlmeyer, R. A. Spotts, M. Serdani, P. W. Crous, K. W. Hughes, K. Matsuura, E. Langer, G. Langer, W. A. Untereiner, R. Lücking, B. Büdel, D. M. Geiser, A. Aptroot, P. Diederich, I. Schmitt, M. Schultz, R. Yahr, D. S. Hibbett, F. Lutzoni, D. J. McLaughlin, J. W. Spatafora & R. Vilgalys. 2006. Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature 443: 818–822.

[K-M02a] Klein-MacPhee, G. 2002a. Smelts. Family Osmeridae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 162–170. Smithsonian Institute Press: Washington.

[K-M02b] Klein-MacPhee, G. 2002b. Pipefishes and seahorses. Family Syngnathidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 321–326. Smithsonian Institute Press: Washington.

[K-MC02] Klein-MacPhee, G., & B. B. Collette. 2002. Eelpouts. Family Zoarcidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 466–474. Smithsonian Institute Press: Washington.

[L91] Liehne, P. F. S. 1991. An Atlas of the Mosquitoes of Western Australia. Health Department of Western Australia.

[MS98] Margulis, L., & K. V. Schwartz. 1998. Five Kingdoms: An Illustrated Guide to the Phyla of Life on Earth 3rd ed. W. H. Freeman and Company: New York.

[M02] Munroe, T. A. 2002. Herrings. Family Clupeidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 111–160. Smithsonian Institute Press: Washington.

[N00] Nichols, G. L. 2000. Food-borne protozoa. British Medical Bulletin 56 (1): 209–235.

[OH02] Olsen, L. E., & M. A. Hoy. 2002. Heat curing Metaseiulus occidentalis (Nesbitt) (Acari, Phytoseiidae) of a fitness-reducing microsporidium. Journal of Invertebrate Pathology 79: 173–178.

[OV95] Ovčarenko, N. A., & I. Vima. 1995. [Issia singulati sp. n. (Microsporidia, Telomyxidae): a new species of Microsporidia from Chironomus singulatus Meigen larvae from Kiev’s water reservoir]. Gidrobiologicheskii Zhurnal 31 (5): 78–83.

[SM03] Saldarriaga, J. F., M. L. McEwan, N. M. Fast, F. J. R. Taylor & P. J. Keeling. 2003. Multiple protein phylogenies show that Oxyrrhis marina and Perkinsus marinus are early branches of the dinoflagellate lineage. International Journal of Systematic and Evolutionary Microbiology 53: 355–365.


Uranotaenia lateralis, copyright Graeme V. Cocks.

Belongs within: Culicidae.

Uranotaenia is a genus of generally very small mosquitoes with reduced micropilosity on the wings (Liehne 1991).

Characters (from Liehne 1991): Adult generally very small; vertex mainly with broad flat scales; proboscis long, slender, frequently swollen apically; palps short; scutum usually very strongly arched; pleura often with distinct lateral lines of iridescent scales; scutellum with broad flat scales; abdomen short, blunt-ended; claws short, simple and subequal, pulvilli absent; wing frequently with some pale iridescent scales, microtrichia of membrane fine and seemingly absent; anal wing vein sharply downcurved at end, ending slightly anterior to base of cubital fork. Larva with head often longer than wide; antenna short; maxillary suture never developed; setae 5-C and 6-C often developed into single flatted spines; lateral comb always present, usually as single row on basal plate; siphon well developed with distinct acus, pecten almost always developed, seta 1-S a single pair and never inserted at base; saddle usually complete.

<==Uranotaenia C09
    |  i. s.: U. lowii C09
    |         U. sapphirina S81
    |--U. (Pseudoficalbia) L91
    |    |--U. (P.) diagonalis L91
    |    `--U. (P.) hirsutifemora L91
    `--U. (Uranotaenia) L91
         |--U. (U.) albescens Taylor 1914 L91
         |--U. (U.) albosternopleura L91
         |--U. (U.) amiensis L91
         |--U. (U.) antennalis L91
         |--U. (U.) argyrotarsis L91
         |--U. (U.) lateralis L91
         |--U. (U.) moresbyensis L91
         |--U. (U.) nivipes L91
         |--U. (U.) novaguinensis L91
         `--U. (U.) tibialis L91

*Type species of generic name indicated


[C09] Chaverri, L. G. 2009. Culicidae (mosquitos, zancudos). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 369–388. NRC Research Press: Ottawa.

[L91] Liehne, P. F. S. 1991. An Atlas of the Mosquitoes of Western Australia. Health Department of Western Australia.

[S81] Stone, A. 1981. Culicidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 341–350. Research Branch, Agriculture Canada.


Tripteroides, probably T. atripes, copyright John Tann.

Belongs within: Culicidae.

Tripteroides is a genus of mosquitoes which possess bristles on the spiracular area but whose wings lack bristles on the stem vein and ventral surface of the subcosta (Liehne 1991).

Characters (from Liehne 1991): Adult with vertex clothed with broad flat scales, upright forked scales few in number and confined to occiput; scutal scaling variable; scutellum with broad flat scales; spiracular bristles present, lower mesepimeral bristles absent; pleura covered with broad flat appressed scales; legs slender, hindtarsus I usually longer than tibia; claws of female equal, simple, pulvilli absent; abdomen blunt-ended, segment VIII of female usually broad and bristly. Larva with head small; antenna short with seta 1-A small, inserted beyond midpoint of antenna; thorax and abdomen generally clothed in stout stellate setae; thoracic spines present, generally well developed; lateral comb always a single row of teeth; seta 1-VIII always more strongly developed than other pentad hairs; pecten absent, siphon covered in numerous setae, some of which may form a 'false' pecten; ventral brush (seta 4-X) always a single pair of setae.

<==Tripteroides L91
    |  i. s.: T. bambusa RK04
    |--T. (Tripteroides) L91
    |    |--T. (T.) magnesianus (Edwards 1924) L91
    |    `--T. (T.) quasiornatus L91
    |--T. (Polylepidomyia) L91
    |    |--T. (P.) atripes (Skuse 1889) [incl. T. apicotriangulata Theobald 1910] L91
    |    `--T. (P.) punctolateralis (Theobald 1903) [incl. T. occidentalis Brug 1934] L91
    `--T. (Rachisoura) L91
         |--T. (R.) sylvestris [incl. T. sylvestris var. hilli] L91
         `--T. (R.) tasmaniensis L91

*Type species of generic name indicated


[L91] Liehne, P. F. S. 1991. An Atlas of the Mosquitoes of Western Australia. Health Department of Western Australia.

[RK04] Russell, R. C., & B. H. Kay. 2004. Medical entomology: changes in the spectrum of mosquito-borne disease in Australia and other vector threats and risks, 1972–2004. Australian Journal of Entomology 43 (3): 271–282.


Gram-stained culture of Bacillus subtilis, copyright Y tambe.

Belongs within: Bacilli.

Bacillus is a diverse genus of rod-shaped, aerobic or facultatively anaerobic Gram-positive bacteria that produce dormant endospores when exposed to unfavorable conditions.

    |--+--B. pumilus VK03
    |  `--B. subtilis VK03 [incl. B. subtilis var. niger CV66]
    `--+--B. firmus VK03
       |--B. lentus VK03
       |--+--B. benzoevorans VK03
       |  `--+--B. circulans VK03
       |     `--B. nealsonii Venkateswaran, Kempf et al. 2003 VK03
       |--+--B. megaterium VK03
       |  `--+--B. flexus VK03
       |     `--B. simplex VK03
       `--+--+--‘B. macroides’ VK03
          |  `--B. psychrosaccharolyticus VK03
          `--+--B. niacini VK03
             `--+--B. fumarioli VK03
                `--‘B. pseudomegaterium’ VK03

Bacillus incertae sedis:
  B. aeolius Gugliandolo, Maugeri et al. 2003VP IJSEM03
  B. alcalophilus PHK96
  B. amyloliquefaciens PHK96
  B. anthracis KE03
  B. azotofixans E92
  B. brevis CM92
  ‘B. caldolyticus’ DC03
  B. caldotenax C-S02
  B. cereus VK03
  B. coagulans PHK96
  B. fastidiosus PHK96
  B. haloalkaliphilus MO03
  B. halodenitrificans SJ04
  B. halodurans Z03
  B. infernus KGP02
  B. larvae White 1906 BB37
  B. laterosporus Z92
  B. licheniformis VK03
  B. methanolicus DC03
  B. mycoides VK03
  B. nitritollens Z92
  B. pantothenticus Z92
  B. pasteurii PHK96
  B. pluton White 1906 BB37
  B. popilliae PHK96
  B. psychrophilus PHK96
  B. pulvefaciens Z92
  B. pycnoticus AS92
  B. schlegelii AS92
  B. selenitireducens Switzer Blum et al. 2001 JC08
  B. sphaericus VK03
  B. stearothermophilus MN03
  B. subterraneus Kanso, Greene & Patel 2002 KGP02
  B. thermantarcticus Nicolaus et al. 2002 JC08
  B. thermocatenulatus DC03
  B. thermocloacae DC03
  B. thermoruber DC03
  B. thuringiensis L91
    |--B. t. var. thuringiensis L91
    `--B. t. var. israelensis L91

*Type species of generic name indicated


[AS92] Aragno, M., & H. G. Schlegel. 1992. The mesophilic hydrogen-oxidizing (knallgas) bacteria. In: Balows, A., H. G. Trüper, M. Dworkin, W. Harder & K.-H. Schleifer (eds) The Prokaryotes: A handbook on the biology of bacteria: Ecophysiology, isolation, identification, applications 2nd ed. vol. 1 pp. 344–384. Springer-Verlag: New York.

[BB37] Bishopp, F. C., & C. E. Burnside. 1937. In memorian: Gershom Franklin White. Proceedings of the Entomological Society of Washington 39 (7): 184–188.

[C-S02] Cavalier-Smith, T. 2002. The neomuran origin of archaebacteria, the negibacterial root of the universal tree and bacterial megaclassification. International Journal of Systematic and Evolutionary Microbiology 52: 7–76.

[CM92] Coughlan, M. P., & F. Mayer. 1992. The cellulose-decomposing bacteria and their enzyme systems. In: Balows, A., H. G. Trüper, M. Dworkin, W. Harder & K.-H. Schleifer (eds) The Prokaryotes: A handbook on the biology of bacteria: Ecophysiology, isolation, identification, applications 2nd ed. vol. 1 pp. 460–516. Springer-Verlag: New York.

[CV66] Curds, C. R., & J. M. Vandyke. 1966. The feeding habits and growth rates of some freshwater ciliates found in activated-sludge plants. Journal of Applied Ecology 3: 127–137.

[DC03] Duan, Y. P., H. F. Castro, T. E. Hewlett, J. H. White & A. V. Ogram. 2003. Detection and characterization of Pasteuria 16S rRNA gene sequences from nematodes and soils. International Journal of Systematic and Evolutionary Microbiology 53: 105–112.

[E92] Eady, R. R. 1992. The dinitrogen-fixing bacteria. In: Balows, A., H. G. Trüper, M. Dworkin, W. Harder & K.-H. Schleifer (eds) The Prokaryotes: A handbook on the biology of bacteria: Ecophysiology, isolation, identification, applications 2nd ed. vol. 1 pp. 534–553. Springer-Verlag: New York.

[IJSEM03] IJSEM. 2003. Validation list no. 94. Validation of publication of new names and new combinations previously effectively published outside the IJSEM. International Journal of Systematic and Evolutionary Microbiology 53: 1701–1702.

[JC08] Judicial Commission of the International Committee on Systematics of Prokaryotes. 2008. Status of strains that contravene Rules 27 (3) and 30 of the International Code of Nomenclature of Bacteria. Opinion 81. International Journal of Systematic and Evolutionary Microbiology 58: 1755–1763.

[KGP02] Kanso, S., A. C. Greene & B. K. C. Patel. 2002. Bacillus subterraneus sp. nov., an iron- and manganese-reducing bacterium from a deep subsurface Australian thermal aquifer. International Journal of Systematic and Evolutionary Microbiology 52: 869–874.

[KE03] Kobayashi, K., S. D. Ehrlich, A. Albertini, G. Amati, K. K. Andersen, M. Arnaud, K. Asai, S. Ashikaga, S. Aymerich, P. Bessieres, F. Boland, S. C. Brignell, S. Bron, K. Bunai, J. Chapuis, L. C. Christiansen, A. Danchin, M. Débarbouillé, E. Dervyn, E. Deuerling, K. Devine, S. K. Devine, O. Dreesen, J. Errington, S. Fillinger, S. J. Foster, Y. Fujita, A. Galizzi, R. Gardan, C. Eschevins, T. Fukushima, K. Haga, C. R. Harwood, M. Hecker, D. Hosoya, M. F. Hullo, H. Kakeshita, D. Karamata, Y. Kasahara, F. Kawamura, K. Koga, P. Koski, R. Kuwana, D. Imamura, M. Ishimaru, S. Ishikawa, I. Ishio, D. Le Coq, A. Masson, C. Mauël, R. Meima, R. P. Mellado, A. Moir, S. Moriya, E. Nagakawa, H. Nanamiya, S. Nakai, P. Nygaard, M. Ogura, T. Ohanan, M. O’Reilly, M. O’Rourke, Z. Pragai, H. M. Pooley, G. Rapoport, J. P. Rawlins, L. A. Rivas, C. Rivolta, A. Sadaie, Y. Sadaie, M. Sarvas, T. Sato, H. H. Saxild, E. Scanlan, W. Schumann, J. F. M. L. Seegers, J. Sekiguchi, A. Sekowska, S. J. Séror, M. Simon, P. Stragier, R. Studer, H. Takamatsu, T. Tanaka, M. Takeuchi, H. B. Thomaides, V. Vagner, J. M. van Dijl, K. Watabe, A. Wipat, H. Yamamoto, M. Yamamoto, Y. Yamamoto, K. Yamane, K. Yata, K. Yoshida, H. Yoshikawa, U. Zuber & N. Ogasawara. 2003. Essential Bacillus subtilis genes. Proceedings of the National Academy of Sciences of the USA 100 (8): 4678–4683.

[L91] Liehne, P. F. S. 1991. An Atlas of the Mosquitoes of Western Australia. Health Department of Western Australia.

[MN03] Moffitt, M. C., & B. A. Neilan. 2003. Evolutionary affiliations within the superfamily of ketosynthases reflect complex pathway associations. Journal of Molecular Evolution 56: 446–457.

[MO03] Müller, V., & A. Oren. 2003. Metabolism of chloride in halophilic prokaryotes. Extremophiles 7: 261–266.

[PHK96] Prescott, L. M., J. P. Harley & D. A. Klein. 1996. Microbiology 3rd ed. Wm. C. Brown Publishers: Dubuque (Iowa).

[SJ04] Spring, S., U. Jäckel, M. Wagner & P. Kämpfer. 2004. Ottowia thiooxydans gen. nov., sp. nov., a novel facultatively anaerobic, N2O-producing bacterium isolated from activated sludge, and transfer of Aquaspirillum gracile to Hylemonella gracilis gen. nov., comb. nov. International Journal of Systematic and Evolutionary Microbiology 54: 99–106.

[VK03] Venkateswaran, K., M. Kempf, F. Chen, M. Satomi, W. Nicholson & R. Kern. 2003. Bacillus nealsonii sp. nov., isolsted from a spacecraft-assembly facility, whose spores are γ-radiation resistant. International Journal of Systematic and Evolutionary Microbiology 53: 165–172.

[Z03] Zeigler, D. R. 2003. Gene sequences useful for predicting relatedness of whole genomes in bacteria. International Journal of Systematic and Evolutionary Microbiology 53: 1893–1900.

[Z92] Zumpft, W. G. 1992. The denitrifying prokaryotes. In: Balows, A., H. G. Trüper, M. Dworkin, W. Harder & K.-H. Schleifer (eds) The Prokaryotes: A handbook on the biology of bacteria: Ecophysiology, isolation, identification, applications 2nd ed. vol. 1 pp. 554–582. Springer-Verlag: New York.


Devil's Hole pupfishes Cyprinodon diabolis, photographed by Olin Feuerbacher.

Belongs within: Atherinomorpha.
Contains: Fundulidae, Poeciliidae.

The Cyprinodontiformes, killifishes and related forms, are a cosmopolitan group of small fishes usually found in estuarine or marine environments (Bond 1996).

Characters (from Bond 1996). Soft-rayed physoclists with abdominal pelvic fins. Single dorsal fin set behind middle of body. Caudal fin with varying number of rays, never forked. Orbitosphenoid, mesocoracoid, and basisphenoid absent; maxillaries excluded from border of mouth by premaxillaries. Lateral line canal system incomplete or not well formed on head and body.

Cyprinodontiformes [Microcyprini]
    |  i. s.: Aplocheilichthys MWW89
    |           |--A. johnstoni CS77
    |           `--A. spilauchen MWW89
    |--Aplocheilidae [Aplocheilinae, Aplocheiloidei] B-RB13
    |    |--Fundulopanchax B96
    |    |--Aphiosemion B96
    |    |--Epiplatys B96
    |    |--Pachypanchax playfairii B96, ND13
    |    `--Aplocheilus L91
    |         |--A. latipes L91
    |         `--A. panchax CS77
    `--Cyprinodontoidei B-RB13
         |  i. s.: Profundulus B96 [Profundulidae B-RB13]
         |           `--P. hildebrandi B96
         |         Valencia B96 [Valenciidae B-RB13]
         |           `--V. hispanica B96
         |         Anablepidae B-RB13
         |           |--Jenynsia [Jenynsiidae] B96
         |           |    `--J. multidentata (Jenyns 1842) MGV06
         |           `--Anableps LD09
         |                |--A. anableps (Linnaeus 1758) LD09 [incl. A. tetrophthalmus Günther 1866 F16]
         |                `--A. microleps Müller & Troschel 1844 PC01
         |         Goodeidae B-RB13
         |           |  i. s.: Tapatia occidentalis G88
         |           |--Goodeinae B96
         |           `--Empetrichthyinae B96
         |                |--Crenichthys B96
         |                `--Empetrichthys B96
         |                     |--E. erdisi G88
         |                     `--E. latos USDI77
         |         Nothobranchius B96 [Nothobranchiidae B-RB13]
         |           `--N. rachowi B96
         |         Rivulidae B-RB13
         |           |--Pterolebias B96
         |           |--Cynolebias B96
         |           `--Rivulus R88
         |                |--R. cylindraceus R88
         |                |--R. garciai R88
         |                |--R. harti [=Haplochilus harti; incl. R. micropus Günther 1866] F16
         |                |--R. heyei R88
         |                |--R. insulaepinorum R88
         |                |--R. isthmensis F15
         |                |--R. marmoratus R86
         |                `--R. roloffi R88
         |--+--Fundulidae ND13
         |  `--Poeciliidae ND13
         `--Cyprinodontidae A02
              |--Kosswigichthys B96
              |--Jordanella floridae B96, ND13
              |--Floridichthys carpio B96, R86
              |--Prolebias G88
              |--Cubanichthys B96
              |    |--C. cubensis R88
              |    `--C. pengelleyi R88
              |--Aphanius CA00
              |    |--A. dispar (Rüppell 1829) CA00
              |    |--A. ginaonis (Holly 1929) CA00
              |    |--A. mento (Heckel 1843) CA00
              |    |--A. persicus (Jenkins 1910) CA00
              |    |--A. sophiae (Heckel 1849) CA00
              |    `--A. vladykovi Coad 1988 CA00
              `--Cyprinodon ND13
                   |--C. atrorus S92
                   |--C. beltrani HB81
                   |--C. bifasciatus S92
                   |--C. bondi R88
                   |--C. diabolis B96
                   |--C. dispar CS77
                   |--C. elegans USDI77
                   |--C. laciniatus R88
                   |--C. maya HB81
                   |--C. nevadensis USDI77
                   |    |--C. n. nevadensis USDI77
                   |    |--C. n. calidae USDI77
                   |    `--C. n. pectoralis USDI77
                   |--C. pecosensis B96
                   |--C. radiosis B96
                   `--C. variegatus Lacépède 1803 D83 [incl. C. baconi R86, C. dearborni R86, C. jamaicensis R86]

*Type species of generic name indicated


[A02] Able, K. W. 2002. Killifishes. Family Fundulidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 292–297. Smithsonian Institute Press: Washington.

[B-RB13] Betancur-R., R., R. E. Broughton, E. O. Wiley, K. Carpenter, A. López, C. Li, N. I. Holcroft, D. Arcila, M. Sanciangco, J. C. Cureton, II, F. Zhang, T. Buser, M. A. Campbell, J. A. Ballesteros, A. Roa-Varon, S. Willis, W. C. Borden, T. Rowley, P. C. Reneau, D. J. Hough, G. Lu, T. Grande, G. Arratia & G. Ortí. 2013. The tree of life and a new classification of bony fishes. PLoS Currents Tree of Life April 18 2013. doi: 10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288.

[B96] Bond, C. E. 1996. Biology of Fishes 2nd ed. Saunders College Publishing: Fort Worth.

[CA00] Coad, B. W., & A. Abdoli. 2000. Systematics of an isolated population of tooth-carp from northern Iran (Actinopterygii: Cyprinodontidae). Zoology in the Middle East 21: 87–102.

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[D83] Darovec, J. E. 1983. Sciaenid fishes (Osteichthyes: Perciformes) of western penisular Florida. Memoirs of the Hourglass Cruises 6 (3): 1–73.

[F15] Fowler, H. W. 1915. Cold-blooded vertebrates from Florida, the West Indies, Costa Rica, and eastern Brazil. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (2): 243–269.

[F16] Fowler, H. W. 1916. The fishes of Trinidad, Grenada, and St. Lucia, British West Indies. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (3): 520–548.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[HB81] Humphries, J. M., F. L. Bookstein, B. Chernoff, G. R. Smith, R. L. Elder & S. G. Poss. 1981. Multivariate discrimination by shape in relation to size. Systematic Zoology 30 (3): 291–308.

[LD09] Li, B., A. Dettaï, C. Cruaud, A. Couloux, M. Desoutter-Meniger & G. Lecointre. 2009. RNF213, a new nuclear marker for acanthomorph phylogeny. Molecular Phylogenetics and Evolution 50: 345–363.

[L91] Liehne, P. F. S. 1991. An Atlas of the Mosquitoes of Western Australia. Health Department of Western Australia.

[MGV06] Mai, A. C. G., A. M. Garcia & J. P. Vieira. 2006. Ecologia alimentar do barrigudinho Jenynsia multidentata (Jenyns, 1842) (Pisces: Cyprinodontiformes) no estuário da Laguna dos Patos, Rio Grande do Sul, Brasil. Comunicações do Museo de Ciências e Tecnologia da PUCRS, Serie Zoologia 19 (1): 3–18.

[MWW89] Miller, P. J., J. Wright & P. Wongrat. 1989. An Indo-Pacific goby (Teleostei: Gobioidei) from West Africa, with systematic notes on Butis and related eleotridine genera. Journal of Natural History 23: 311–324.

[ND13] Near, T. J., A. Dornburg, R. I. Eytan, B. P. Keck, W. L. Smith, K. L. Kuhn, J. A. Moore, S. A. Price, F. T. Burbrink, M. Friedman & P. C. Wainwright. 2013. Phylogeny and tempo of diversification in the superradiation of spiny-rayed fishes. Proceedings of the National Academy of Sciences of the USA 110 (31): 12738–12743.

[PC01] Pichelin, S., & T. H. Cribb. 2001. The status of the Diplosentidae (Acanthocephala: Palaeacanthocephala) and a new family of acanthocephalans from Australian wrasses (Pisces: Labridae). Folia Parasitologica 48: 289–303.

[R88] Rauchenberger, M. 1988. Historical biogeography of poeciliid fishes in the Caribbean. Systematic Zoology 37 (4): 356–365.

[R86] Rivas, L. R. 1986. Comments on Briggs (1984): freshwater fishes and biogeography of Central America and the Antilles. Systematic Zoology 35 (4): 633–639.

[S92] Smith, G. R. 1992. Introgression in fishes: significance for paleontology, cladistics, and evolutionary rates. Systematic Biology 41 (1): 41–57.

[USDI77] United States Department of the Interior. 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.