Apogonini Günther 1859

Mabuchi, Kohji, Fraser, Thomas H., Song, Hayeun, Azuma, Yoichiro & Nishida, Mutsumi, 2014, Revision of the systematics of the cardinalfishes (Percomorpha: Apogonidae) based on molecular analyses and comparative reevaluation of morphological characters, Zootaxa 3846 (2), pp. 151-203 : 178-179

Tribe Apogonini Günther 1859

Type genus Apogon Lacepède 1801

Diagnosis. Members of the Apogoninae : dorsal fin VI+I,9; anal fin II,8; developed gill rakers 9–19; posttemporal edge smooth or weakly serrate; anterior nare with low rim and flap; preopercle smooth on ridge, serrate to smooth on vertical edge, horizontal edge an unossified large flap; head and body with ctenoid or cycloid scales; pored lateral-line scales 23–25, scales with one pore above center and one below central pore; intestine and stomach pale (except two species); head and body reddish with or without blackish markings or head and body brownish without bars or stripes, both color patterns may be translucent in some species.

Other characteristics. one supernumerary spine; branched first segmented dorsal and anal ray; ctenoid or cycloid scales on predorsal, cheek, breast, two pelvic scales, and body; cycloid scale on opercle and onto base of caudal fin; pored lateral-line scales simple with one pore on upper side and one on lower side; pectoral fin-rays 11–16; 0–3 supraneurals; pored lateral-line scales from posttemporal to base of caudal fin; caudal fin forked or rounded; 9+8 segmented principal caudal rays, 15 branched, upper and lower unbranched; unbranched procurrent rays, longest segmented; teeth on premaxilla, dentary, vomer, palatine, all villiform (except one species canine-like on premaxilla and dentary) or absent on palatine; six infraorbitals, bony shelf on third infraorbital; supramaxilla absent; basisphenoid present, reduced or absent; anterior ceratohyal notched; 10+14 vertebrae; 8 ribs; 8–9 epineurals; uroneurals absent; three epurals; five free hypurals, 1–2 fused and 3–4 fused, 1–2 fused and 3–4 fused to terminal centrum; free parhypural; low crest on PU2.

Distribution. Apogon sensu stricto is found in all tropical regions, Zapogon Fraser 1972 in the Atlantic and Indo-Pacific, Phaeoptyx Fraser & Robins 1970 and Paroncheilus Smith 1964 in the Eastern and Western Atlantic Ocean and Astrapogon Fowler 1907 confined to the Western Atlantic Ocean.

Remarks. This tribe contains five genera, Astrapogon , Apogon sensu stricto, Phaeoptyx , Paroncheilus and Zapogon , corresponding to the clade II in the molecular trees ( Figs. 2–6 , Table 4 ). Members have a large, unossified preopercular flap (a proposed synapomorphy), simple pore arrangement on lateral-line scales, six first dorsal spines and color patterns. Monophyly of Apogon sensu stricto (H10) was, however, statistically rejected by AU test ( Table 6 ). In the molecular trees, Apogon sensu stricto was divided into three ( Apogon -1, 2, and 3) or two ( Apogon -1 and 2+3) lineages. One of the two or three lineages, Apogon -1, was sister to the clade including all the other members of this tribe. This basal dichotomy agrees with geographic distributions: species of the former clade ( Apogon -1) distributed in Indo-Pacific Basin and those of the latter clade basically in Atlantic Basin with a small group of Apogon -3 within it occurring in Eastern Pacific. These molecular results suggest that species of Apogonini have been firstly separated between the Atlantic and Indo-Pacific regions and then an Atlantic species invaded to Eastern Pacific. Asperapogon Smith 1961 is an available name for some or all of the Indo-Pacific species ( Apogon -1) as a genus or subgenus. A morphological diagnosis for Asperapogon awaits determination of species composition. Type species of Apogon is Apogon ruber Lacepède 1801 , a synonym of A. imberbis ( Linnaeus 1758) from the Eastern Atlantic Basin and the Mediterranean Sea. No other subfamilies or tribes occur in the Atlantic Basin or Eastern Pacific.

Figures

FIGURE 2. Phylogenetic tree from the partitioned maximum-likelihood (ML) analysis. Numbers besides internal branches indicate bootstrap probabilities (BPs) from 1000 replicates (only those of> 50% shown). Species names of recent Ostorhinchus [we included here Apogonichthyoides melas as a possible Ostorhinchus, following Mabuchi et al. (2006)] were blue, those of Apogon sensu lato [genus Apogon in Fraser (1972)] excepting the Ostorhinchus red, and those of genus Rhabdamia in Fraser (1972) green. The names of the twelve major clades (clades I to XII) within the Apogoninae were indicated on the left of the corresponding clades. Four species, which phylogenetic positions were largely different among the ML, Bayesian (BA: Fig. 3), and most parsimonious (MP: Figs. 4 and 5) tress, were indicated by arrows with asterisks.

FIGURE 3. Phylogenetic tree (50% majority-rule consensus tree) from the partitioned Bayesian (BA) analysis. Numbers besides internal branches indicate Bayesian posterior probabilities (PPs: shown as percentages, only those of> 50% shown). Species names were colored as in Figure 2. The names of the twelve major clades were indicated as in Figure 2. Four species, which phylogenetic positions were largely different among the ML (Fig. 2), BA (Fig. 3), and most parsimonious (MP: Figs. 4 and 5) tress, were indicated by arrows with asterisks.

FIGURE 4. Strict consensus tree of two of the five most parsimonious (MP) trees (MP tree-A). Numbers besides internal branches indicate bootstrap probabilities (BPs) from 1000 replicates (only those of> 50% shown). Topological incongruity between the two MP trees denoted by an arrowhead. Four species, which phylogenetic positions were largely different among the ML (Fig. 2), BA (Fig. 3), and MP (Figs. 4 and 5) tress, were indicated by arrows with asterisks. Species names were colored as in Figure 2. The names of the twelve clades were indicated as in Figure 2.

FIGURE 5. Strict consensus tree of the remaining three of the five most parsimonious (MP) trees (MP tree-B). Topological incongruities between the three MP trees denoted by arrowheads. Four species, which phylogenetic positions were largely different among the ML (Fig. 2), BA (Fig. 3), and MP (Figs. 4 and 5) tress, were indicated by arrows with asterisks. This tree was largely different from the MP tree-A (Fig. 4) in the positions of Pterapogon kauderni and Vincentia novaehollandiae.

FIGURE 6. Strict consensus tree of the ML, BA and MP trees shown in Figures 2-5. Species names were colored as in Figures 2–5. The names of the twelve major clades (clades I to XII) within the Apogoninae were indicated besides the corresponding clades, while the names of the thirteen tribes (defined based on morphological characters) on the right of corresponding major and minor clades. The tribe Lepidamiini is not in the tree. Tribes with solid bars include species of Apogon sensu lato, and those with open bars do not include it. Geographic ranges were indicated on the right of the clades only for the species of clade II (Apogonini), all the other apogonids occurring on Indo-Pacific Basin. For the revised genus names of some species, see Appendix A.