Danionin

The danionins are a group of small minnow-type fish belonging to the family Cyprinidae. Members of this group are mostly in the genera Danio, Devario and Rasbora. They are primarily native to the fresh waters of South and Southeast Asia, with fewer species in Africa. Many species are brightly coloured and are available as aquarium fish worldwide. Danio species tend to have horizontal stripes, rows of spots, or vertical bars, and often have long barbels. Devario species tend to have vertical or horizontal bars, and short rudimentary barbels, if barbels are present at all. All danionins are egg scatterers and breed in the rainy season in the wild. They are carnivores living on insects and small crustaceans.

Danionins
Danio rerio — zebra danio
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Order:	Cypriniformes
Family:	Cyprinidae
Subfamily:	Danioninae Bleeker 1863[1]
Genera
Amblypharyngodon Aspidoparia Barilius Betadevario Boraras Cabdio Chela Chelaethiops Danio Danionella Devario Engraulicypris Esomus Fangfangia Horadandia Inlecypris Laubuca Leptocypris Luciosoma Malayochela Microdevario Microrasbora Nematabramis Neobola Opsaridium Opsarius Paedocypris Pectenocypris Raiamas Rasbora Rasboroides Rastrineobola Salmostoma Securicula Sundadanio[2] Trigonopoma Trigonostigma

Taxonomy

The grouping of fish now deemed danionins has been the subject of constant research and speculation throughout the 20th century. Nearly all the fish classed within the genera Danio and Devario were originally placed in the genus Danio upon discovery. However, in the first part of the 20th century, George S. Myers split them into three genera, Danio, Brachydanio, and Daniops. The sole species within Myers' Daniops, D. myersi, has long ago been found to be a synonym of Devario laoensis, but his genus Brachydanio lasted for much longer, as it included most of the fish now classed as Danio, whereas Danio included most of the fish now classed as Devario.[3]

However, Danio dangila and Danio feegradei, both of which had most of the characteristics of the Brachydanio (with the exception that they were much larger than Brachydanio species) were placed within Danios. (Due to this and other misplacing, both Danio and Brachydanio were found to be paraphyletic by Fang Fang in 2003.[4]). In 1941, H.M. Smith attempted to unite all the Brachydanios and Danios species into one genus on the basis of a fish from Thailand which was supposed to bridge the gap. He downgraded both Danio and Brachydanio into subgenera and erected a new subgenus of Allodanio with one member, Allodanio ponticulus, but Myers later pointed out that A. ponticulus was actually a member of the genus Barilius.[3]

The danionin group was thought to include Parabarilius, Danio, Brachydanio, and Danionella. In this scheme, danionins were distinguished from other cyprinids by the uniquely shared character of the "danionin notch", a large and peculiarly shaped indentation in the medial margin of the mandibles; this feature is not noted in rasborins, esomins, bariliins, or chelins. However, all of these categories at that time were informal. Microrasbora was not considered to be a part of the danionins, nor even closely related to Danionella, a part of the danionins as understood at that time.[5]

In the late 1980s and 1990s, doubts grew about the validity of Brachydanio, with species being referred to their original naming of Danio, and Fang Fang determined that the genus Danio, recognized up to that point, was paraphyletic. Fang restricted Danio to the species in the "D. dangila species group", which at the time comprised nine species including D. dangila, D. rerio, D. nigrofasciatus, and D. albolineatus; the remaining Danio species were moved to Devario, which at this time included D. malabaricus, D. kakhienensis, D. devario, D. chrysotaeniatus, D. maetaengensis, D. interruptus, and D. apogon.

The only Danio species to have been consistently called Danio were D. dangila and D. feegradei. As D. dangila was the first discovered Danio (or type) the name Danio had to remain with D. dangila, which is why the vast majority of species were moved to Devario.

Also, the sister group to Devario was deemed to be a clade formed by Inlecypris and Chela, and more controversially, Esomus was found to be the sister group of Danio. The relationships of Sundadanio, Danionella, and Microrasbora remained unresolved. The danionin notch was found to not supported to be a danionin synapomorphy.[4]

In another paper, Celestichthys margaritatus was described as a new species of the Danioninae. Apparently, it is most closely related to Microrasbora erythromicron; the other Microrasbora species differ significantly from Celestichthys. The genus is identified as a danionin due specializations of its lower jaw and its numerous anal fin rays. Though it lacks a danionin notch, Celestichthys exhibits the "danionin mandibular knob", a bony process on the side of the mandible behind the danionin notch or where the notch should be; it is perhaps diagnostic of danionins. This knob is better developed in males than females. The fish of Rasborinae almost invariably have anal fins with three spines and five rays. Celestichthys has three anal fin spines and 8-10 anal fin rays. Also, rasborins have the generalized cyprinid principal caudal fin ray count of 10/9, while all Asian cyprinids with fewer than 10/9 principal caudal fin rays are all diminutive species of Danioninae, including Celestichthys, M. erythromicron, Danionella, and Paedocypris.[6]

In 2007, an analysis of the phylogenetic relationships of the recently described genus Paedocypris was published, placing it as the sister taxon to Sundadanio. The clade formed by these two genera was found to be sister to a clade including many danionin genera, as well as some rasborin genera such as Rasbora, Trigonostigma, and Boraras, making the danionin group paraphyletic without these rasborin genera based on these results. This paper considered the danionin genera to be within a larger Rasborinae.[7]

Also in 2007, another study analyzed the relationships of Danio. These authors considered Rasborinae to have priority over Danioninae, suggesting that they have the same meaning. Also, Danio was found to be the sister group of a clade including Chela, Microrasbora, Devario, and Inlecypris, rather than in a clade exclusively with either Devario or Esomus as in previous studies. This paper supported the close relationship of "Microrasbora" erythromicron to Danio species; however, this study did not include Celestichthys, which was noted by Roberts as being likely to include Erythromicron, but with further research needed.[6][8]

Tanichthys is often regarded as a danionin by aquarists and grouped as such in some older aquatic publications, but no scientific basis exists for this, a fact stated on numerous occasions by Brittan and others.[3] It is more closely related to the Rasbora species. The danionins can be classed as a subfamily Danioninae which is increasingly gaining credibility as a subfamily distinct from the Rasboriniae within the family Cyprinidae.[9] However, in Nelson, 2006, Danioninae was listed as a synonym of Rasborinae.[10] However, neither inter- nor intrarelationships among the "rasborins" has as yet been thoroughly analyzed.[4]

A number of the species have only been recently discovered, in remote inland areas of Laos and Myanmar, and do not yet have scientific names. They are listed as Danio or Devario sp "xxxx" within the relevant genera and disambiguation pages.

In the aquarium

They are generally active swimmers, occupying the top half of a tank and eat just about any type of aquarium food. They will not, however, generally eat plants or algae.

Although boisterous and liable to chase each other and other fish, they are good community fish and do not generally attack each other or other fish, although they occasionally nip fins, and like most fish, eat eggs and any fish small enough to fit into their mouths.

These fish are easily stressed by flowing water and bright light. They occur in stagnant water with PH between 3 and 5 caused by peat which accumulates from a dense canopy. It is unknown how the peat and tannins interact with the fish.

Generally, this also results in them being subtropical with temperatures of 20 to 22 °C (68 to 72 °F) often being fine; they are good jumpers, so a tight-fitting lid is recommended.

Common names given to danionin species

Since 2004, many new danionins have been discovered which do not yet have scientific names and many other species, previously known only to the scientific fraternity, have become available in aquarist shops. This has predictably led to total confusion as to the naming of some fish, with some species having up to five different common names in use and some common names bein used for up to four different species. As a result, all danionin common names known to be in use are listed on a separate page:

• List of danionin species by common name

Danionin species scientific names

Individual danionin species are listed within the relevant pages for each genus, but many danionin species have been changed into different genera over the last decades, in some cases repeatedly; similarly, some species have been synonymised with other species and in some cases later unsynonymised, all of which has caused confusion. As a result, a separate page has been created which lists all danionin species and also lists defunct species which have since been synonymised or renamed.

• Danionin species disambiguation

External links

• "Danios and Devarios". Danios and devarios website. Retrieved October 1, 2005.

References

1. Van der Laan, R.; Eschmeyer, W. N.; Fricke, R. (11 November 2014). "Family-group names of Recent fishes". Zootaxa Monograph. 3882: 1–230. doi:10.11646/zootaxa.3882.1.1. PMID 25543675.
2. Tang, KL; Agnew, MK; Hirt, MV; Sado, T; Schneider, LM; Freyhof, J; Sulaiman, Z; Swartz, E; Vidthayanon, C; Miya, M; Saitoh, K; Simons, AM; Wood, RM; Mayden, RL (2010). "Systematics of the subfamily Danioninae (Teleostei: Cypriniformes: Cyprinidae)". Mol Phylogenet Evol. 57 (1): 189–214. doi:10.1016/j.ympev.2010.05.021. PMID 20553898.
3. Brittan, Martin Ralph (1958). "Danios, A guide to the identification, care and breeding of fishes of the genera Danio and Brachydanio". T.F.H.Publications. ASIN B00MFY4BTA. Cite journal requires |journal= (help)
4. Fang, Fang; Douglas, M. E. (2003). Douglas, M. E. (ed.). "Phylogenetic Analysis of the Asian Cyprinid Genus Danio (Teleostei, Cyprinidae)". Copeia. 2003 (4): 714–728. doi:10.1643/IA03-131.1.
5. Roberts, Tyson R. (1986). "Danionella translucida, a new genus and species of cyprinid fish from Burma, one of the smallest living vertebrates". Environmental Biology of Fishes. 16 (4): 231–241. doi:10.1007/BF00842977.
6. Roberts, T. R. (2007). "The "Celestial pearl danio", a new genus and species of colourful minute cyprinid fish from Myanmar (Pisces: Cypriniformes)" (PDF). The Raffles Bulletin of Zoology. 55 (1): 131–140. Retrieved 2009-06-24.
7. Rüber, Lukas; Kottelat, Maurice; Tan, Heok Hui; Ng, Peter KL; Britz, Ralf (2007). "Evolution of miniaturization and the phylogenetic position of Paedocypris, comprising the world's smallest vertebrate". BMC Evolutionary Biology. 7: 1–10. doi:10.1186/1471-2148-7-38. PMC 1838906. PMID 17355618.
8. Mayden, Richard L.; Tang, Kevin L.; Conway, Kevin W.; Freyhof, Jörg; Chamberlain, Sarah; Haskins, Miranda; Schneider, Leah; Sudkamp, Mitchell; Wood Robert M.; Agnew, Mary; Bufalino, Angelo; Sulaiman, Zohrah; Miya, Masaki; Saitoh, Kenji; He, Shunping (2007). "Phylogenetic relationships of Danio within the order Cypriniformes: a framework for comparative and evolutionary studies of a model species". J. Exp. Zool. (Mol. Dev. Evol.). 308B (1): 1–13. doi:10.1002/jez.b.21130. PMID 17171697.
9. Clarke, Matt. "A fishkeeper's guide to danios and devarios". Practical Fishkeeping (March 2005). Archived from the original on 2005-12-24.
10. Nelson, Joseph S. (2006). Fishes of the World. John Wiley & Sons, Inc. ISBN 978-0-471-25031-9.