Phylogenetic Analysis of the Asian Cyprinid Genus Danio (Teleostei, Cyprinidae)
A phylogenetic analysis of Danio (sensu lato), based on 38 morphological characters, shows that Danio, as recognized until now, is paraphyletic. Danio is restricted to species previously recognized as the “Danio dangila species group,” including D. dangila, Danio rerio, Danio nigrofasciatus, and Danio albolineatus. Those species share an “A stripe” on the anal-fin rays, an anterior lateral extension ventrally on the dentary, two or more pigment stripes on the caudal-fin rays and greatly enlarged lamellar nasal. Esomus is the sister group of Danio. Remaining Danio (sensu lato) species are referred to Devario, characterized by a short and wide premaxillary ascending process with a minute apophysis contacting the kinethmoid, a short maxillary barbel, a “P stripe” extending onto the median caudal-fin rays, and infraorbital 5 not or only slightly reduced. Devario includes the species Devario malabaricus, Devario kakhienensis, Devario devario, Devario chrysotaeniatus, Devario maetaengensis, Devario interruptus, and Devario apogon. The sister group of Devario is a clade composed of Inlecypris and Chela. The well-known genus Brachydanio becomes a junior synonym of Danio (sensu stricto). The phylogenetic relationships of small Danio-like species Sundadanio axelrodi, Danionella translucida, Danio erythromicron, and Microrasbora rubescens remain unresolved.Abstract

Cladogram of danionin taxa representing strict consensus tree of two equally parsimonious trees. Tree length 97 steps, consistency index (CI) = 0.4639, homoplasy index (HI) = 0.5361, retention index (RI) = 0.7463, and rescaled consistency index (RC) = 0.3462. Character states mapped according to accelerated optimization; black boxes represent derived states, open boxes are independently acquired homoplastic states, and black/white boxes represent reversals from previously derived states. Figures in boxes represent bootstrap values after 1000 replications; figures in ellipses represent Bremer decay indices

(A) Danio erythromicron (NRM 46956, 18.9 mm SL), left maxilla, lateral view. (B) Devario kakhienensis (NRM 34001, 40.8 mm SL), left maxilla, lateral view. (C) Danio aff. dangila (NRM 46966, 38.1 mm SL), left maxilla, lateral view. (D) Danio albolineatus (NRM 47071, 29.7 mm SL), left lower jaw, ventral view. (E) Danio aff. dangila (NRM 46966, 38.1 mm SL), left lower jaw, ventral view. (F) Sundadanio axelrodi (NRM 47436, 18.7 mm SL), right lower jaw, ventral view. (G) Danio aff. dangila (NRM 46966, 38.1 mm SL), right palatine, lateral view. (H) Devario kakhienensis (NRM 34001, 40.8 mm SL), right palatine, lateral view. (I) Danio albolineatus (NRM 33003, 33.3 mm SL), kinethmoid, dorsal view. (J) Devario kakhienensis (NRM 34001, 40.8 mm SL), kinethmoid, dorsal (a) and anterior (b) view

Danio albolineatus (NRM 33003, 33.3 mm SL), the first four vertebrae, right side, lateral view showing median projection on rib of the fourth vertebra

Devario interruptus (NRM 32194, 40.1 mm SL), head region, dorsal view

(A) Devario kakhienensis (NRM 34003, 52.5 mm SL). (B) Danio. aff. dangila (NRM 46966, 38.1 mm SL). (C) Sundadanio axelrodi (NRM 47436, 18.7 mm SL). Left infraorbital series, lateral view, IO1-IO5, infraorbitals 1–5

(A) Devario devario (NRM 46955, 49.4 mm SL). (B) Danio dangila (NRM 46953, 69.0 mm SL). Upper jaw, dorsal view

(A) Devario devario (NRM 46955, 49.4 mm SL). (B) Danio dangila (NRM 46953, 69.0 mm SL). Left hyopalatine arch, lateral view

(A) Devario malabaricus (NRM 24664, 53.9 mm SL). (B) Devario kakhienensis (NRM 34003, 52.5 mm SL). (C) Danio albolineatus (NRM 33003, 33.3 mm SL). Anterior part of neurocranium, dorsal view
Contributor Notes
Department of Vertebrate Zoology, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05, Stockholm, Sweden. fang.fang@nrm.se