Classification

Goodeids have been named after the first known genus, Goodea, which has been described by David Starr Jordan in 1880 in honour of George Brown Goode (13.02.1851-06.09.1896), his colleague and american ichthyologist, who worked most of his time as a museum administrator.

In 1872 Goode became the assistant of Spencer Baird,  secretary of the Smithsonian Institute, and started to run the fish research program of the U.S. Fish Commision and the Smithsonian Institution in 1873. He effectively ran both till 1887. He authored many books and monographs and wrote more than 100 scientific reports and notes, most of them together with his Smithsonian colleague Tarleton Hoffman Bean.

Goode had been a member of the National Academy of Sciences and the American Academy of Arts and Sciences and died of Pneumonia at Lanier Heights, near Washington as a young man at the age of 45 on Sept.6, 1896. His death left a void at the Smithsonian and together with the departure of Tarleton Bean the year before, research at the Smithsonian came to a stillstand. Goode had been universally liked and respected and had been very important for the American research activities concerning North American fish. 

 

Goodeids are charactrized by some synapomorphys:

First 2-7 middle anal radials fused to proximal radials

Dorsal process of maxillaries greatly reduced

Distal arm of premaxilla straight

Articular reduced

 

Prior to the revision of Lynne R. Parenti in 1981, two genera of egglaying fishes from Nevada have been aligned with Cyprinodontidae or placed in their own family, Empetrichthyidae. Meanwhile these genera with two species each and a few subspecies are recognized as a subfamily within Goodeids: Empetrichyinae. They are characterized by the lack of a pelvic skeleton. The genus Empetrichthys includes Empetrichthys latos from the Pahrump Valley in southern Nye County and the extinct Empetrichthys merriami, Crenichthys the two species Crenichthys nevadae and Crenichthys baileyi with 5 subspecies, Cr. b. albivallis, Cr. b. baileyi, Cr. b. grandis, Cr. b. moapae and Cr. b. thermophilus.

About 45 species from the Mexican Mesa Central are classified within the subfamily Goodeinae. In contrary to the Empetrichthyinae, these species are euviviparous with an internal fertilization. Other synapomorphys of this monophyletic group are:

Anterior rays of anal of males shortened and slightly separated from rest of fin by a notch, producing a gonopodium-like pseudophallus

Eggs are small with little yolk

Embryos and newborn young with ribbon-like extensions from anal region associated with nutrition and respiration, termed trophotaeniae

 

Following Webb & Miller (2002) respectivelyCharacodon audax "El Toboso" Dominguez & Doadrio (2004), in this subfamily are recognized several tribes. The first one is the monogenetic tribe Characodontini including the genus Characodon with 3 described species: Characodon audax, Characodon lateralis and Characodon garmani. The latter became extinct in the late 1960s. The remaining two species are living distinct from the other Goodeid genera in the state of Durango, in the northern part of the Mesa Central. Habitats of these fish are located along the Río Mezquital in the Guadiana valley.

A second and well defined tribe, Ilyodontini, is comprised ofXenotaenia resolanae 3 genera, all of them living on the Pacific slope of the Mesa Central in the states Jalisco, Colima, Michoacán, Morelos and México.

The genus Xenotaenia is monotypic. The only species Xenotaenia resolanae seems to be the most original form of this tribe.

Four ground-dwelling species are summarized in the genus Allodontichtys: Allodontichthys hubbsi, polylepis, tamazulae and zonistius.

The remaining genus Ilyodon is - citing Kingston - a taxonomic nightmare. Some trophical varieties and not well defined forms make it nearly impossible to distinguish species. In addition toAllodontichthys tamazulae morphological troubles, we find less genetic variance between two recognized species  - Ilyodon furcidens and Ilyodon whitei  - than within populations of Ilyodon whitei (after Dominguez et al., 2004). A third species, Ilyodon cortesae has been described and is widely recognized. The two described species Ilyodon xantusi and Ilyodon lennoni are treated as varieties of furcidens respectively whitei from most of the scientists, another undescribed form from the Ameca river may likely be a distinct species, Ilyodon sp. "Ameca", sometimes mistakenly named Ilyodon amecae.Ilyodon whitei

The former two tribes are clearly distinguishable from the rest of the Goodeids. The coming species are more related to each other, nevertheless they can be comprised into 3 tribes.

The first one, Goodeini, is made up of two genera.Ataeniobius toweri One of them and monotypic is  Ataeniobius with the only species toweri endemic to the upper Rio Verde drainage in the State San Luis Potosí. Therewith it is the most eastern living species of Goodeids, and one of few inhabiting a river draining to the Atlantic ocean. 

The second genus Goodea with - at the moment - two described species, Goodea atripinnis and Goodea gracilis is taxonomically nearly as difficult as Ilyodon. Several species have been described and seized again. Even the species gracilis isn't recognized by all scientists. However, Goodea gracilis from Querétaro and San Luis Potosí is treated as endangered, but the remaining atripinnis is definitely the most successful species among Goodeids, inhabiting nearly all areas and habitats of the Mesa Central. The reason for this success is unknown yet.Goodea atripinnis Recent phylogenetical results (Domínguez, pers.comm. 2012) place Ataeniobius within the tribe Girardinichthyini and Goodea within the tribe Chapalichthyini, seizing the tribe Goodeini.

The next tribe Chapalichthyini is containing 7 genera, one of them undescribed. The monotypic genus AmecaAmeca splendenswith the popular Butterfly Splitfin, Ameca splendens is endemic to the Río Ameca and its tributary, the Río Teuchitlán. A visually resembling species can be found within the genus Chapalichthys, namely pardalis from a pool at Tocumbo in Michoacán. One up to two further species are recognized, depending on the author. Chapalichthys encaustus is distributed in Jalisco and Michoacán, Chapalichthys peraticus is restricted to the Chapalichthys encaustussmall endorheic basin of the Represa de San Juanico near Cotija in Michoacán. Chapalichthys peraticus is not accepted as valid by many authors, mainly because there are only few differences to pardalis

The only known predator within Goodeids is Alloophorus robustus from the monotypic(?) genus Alloophorus. These fish grow up to 15cm and are widely distributed in Michoacan, but are not common. This species seems to have his closest relatives within the genus Chapalichthys.Alloophorus robustus

The genus Zoogoneticus is standing a little bit isolated within this tribe, comprising 3 small and colourful species from Jalisco, Michoacán and Guanajuato. Zoogoneticus tequila from the Río Teuchitlán system seemed to be extinct in the wild immediately after its discovery in the early 1990s, but had been rediscovered a few years ago. Zoogoneticus purhepechus had been splitted fromZoogoneticus quitzeoensisZoogoneticus quitzeoensis quite recently. These both species are barely distinguishable.

Xenoophorus is one of the monotypic genera of Goodeids, the only species is Xenoophorus captivus from the states of San Luis Potosí and Zacatecas.Xenoophorus captivus

The genus Xenotoca contains one described species, Xenotoca variata and one undescribed form from the Cuitzeo Xenotoca variatabasin.

The species eiseni and melanosoma, traditionally members of the genus Xenotoca belong definitely to a distinct genus. The name proposed by Webb (1998), Xenotichthys, has not been published officially yet, so the genus is termed "Xenotoca" for the moment following Robert Rush Miller, published after his death 2005. Probably the eiseni from Río Etzatlan will have to be described as an own species."Xenotoca" eiseni

 

The last tribe embraces two different groups, forming a monophyletic group either.

The first group within the tribe Girardinichthyini can be called Allotoca-group, formed by 2 genera.

The smaller genus, Neoophorus with the single species regalis is sometimes included in the remaining genus Allotoca. Neoophorus regalis is restricted to a small locality in the vicinity of Los Reyes, Michoacán.Neoophorus regalis

Allotoca is comprised by an uncertain number of species. Whilst all scientists recognize the species Allotoca dugesii, goslinei, maculata and zacapuensis as valid, the species catarinae, diazi and meeki are treated sometimes as only one species: diazi. Dominguez (2004) noted, that the genetic variance between the 3 forms is very low, lower than within some species. However, examinations will have to take place to clear the situation. Additionally one or two undetermined forms are known, at least one of them is considered as an undescribed species. All of the species except dugesii are restricted to small areas in the states Michoacán and Jalisco and are extremely endangered.Allotoca dugesii

The second group could be called Girardinichthys-group. Four genera belong to this group.

The genus Skiffia with 3 described species, Skiffia francesae, lermae and multipunctata inhabits non current habitats in the states Jalisco, Michoacán and Guanajuato. Skiffia lermae is clear distinguishable from multipunctata, Skiffia francesae might be a form of multipunctata. A recently discovered form called Skiffia sp. "Sayula" might be a variety of multipunctata.Skiffia multipunctata

The genus Neotoca has been reestablished by Dominguez in 2004 as a monotypic genus for Neotoca bilineata, former included in Skiffia. This fish seems to be more related to the remaining two genera than to Skiffia.Neotoca bilineata

The last two genera, Girardinichthys and Hubbsina - sometimes treated as one genus with two subgenera - contain two species each. Girardinichthys viviparus was the first known species of Goodeids. First attempts to keep this fish in captivity might date back to 1772! It is restricted to the endorheic basin of Valle de México. The second species Girardinichthys multiradiatus inhabits waterbodies in the states Morelos, Michoacán and México.Girardinichthys viviparus

Hubbsina is comprised of the late described ireneae and the - maybe - extinct turneri. While ireneae can be found in many specimens in the Zacapu lagoon, turneri has not been found for decades now. Since the identification by Alfred Radda and Manfred Meyer in 2003 (as Girardinichthys ireneae), Hubbsina ireneaeit became clear, that a lot of turneri kept in tanks before had been ireneae. However, after all, it may be that both "species" belong only to one, namely turneri, (following some authors) because the differences between both are few and weakly supported. All four species are characterized by their high number of rays in the dorsal fin (Hubbsina) and additionally anal fin (Girardinichthys).

 

The classification of described Goodeids.

 

 

Subfamily                   Tribe                            Genus                         Species

 

Empetrichthyinae

Empetrichthys

Empetrichthys latos concavus (ext.)

Empetrichthys latos latos

Empetrichthys latos pahrump (ext.)

Empetrichthys merriami (ext.)

Crenichthys

Crenichthys baileyi albivallis

Crenichthys baileyi baileyi

Crenichthys baieyi moapae

Crenichthys baileyi grandis

Crenichthys baileyi thermophilus

Crenichthys nevadae

 

Goodeinae

 Characodontini

 Characodon

Characodon audax

Characodon garmani (ext.)

Characodon lateralis

Ilyodontini

Allodontichthys

Allodontichthys hubbsi

Allodontichthys polylepis

Allodontichthys tamazulae

Allodontichthys zonistius

Ilyodon

Ilyodon cortesae

Ilyodon furcidens

Ilyodon whitei

Xenotaenia

Xenotaenia resolanae

Goodeini

Ataeniobius

Ataeniobius toweri

Goodea

Goodea atripinnis

Goodea gracilis

Chapalichthyini

Alloophorus

Alloophorus robustus

Ameca

Ameca splendens

Chapalichthys

Chapalichthys encaustus

Chapalichthys pardalis

Chapalichthys peraticus

Xenoophorus

Xenoophorus captivus

Xenotoca

Xenotoca variata

Xenotoca cf. variata

"Xenotoca"

"Xenotoca" doadrioi

"Xenotoca" eiseni

"Xenotoca" lyonsi

"Xenotoca" melanosoma

"Xenotoca" cf. melanosoma

 Zoogoneticus

Zoogoneticus purhepechus

Zoogoneticus quitzeoensis

Zoogoneticus tequila

Girardinichthyini

Allotoca

Allotoca catarinae

Allotoca diazi

Allotoca dugesii

Allotoca goslinei

Allotoca maculata

Allotoca meeki

Allotoca zacapuensis

Girardinichthys

Girardinichthys multiradiatus

Girardinichthys viviparus

Hubbsina

Hubbsina ireneae

Hubbsina turneri (ext?)

Neoophorus

Neeophorus regalis

Neotoca

Neotoca bilineata

Skiffia

Skiffia francesae

Skiffia lermae

Skiffia multipunctata