Pseudamiinae Smith 1954

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 : 177-178

Subfamily Pseudamiinae Smith 1954

Type genus Pseudamia Bleeker 1865

Diagnosis. Two separate dorsal fins, VII dorsal spines as VI+I,8–9; anal fin II,8–10; wide separation between dorsal fins; internal support of dorsal spines by serial proximal-middle radials not in close articulation, far apart at 6th between and 7th elements; one supernumerary dorsal spine; 1–2 supraneurals; first anal proximal-middle radial straight; 9+8 principal caudal fin-rays, 15 branched, upper and lower unbranched, caudal fin rounded or rhomboid, mid-line fin-ray longest; supramaxilla and basisphenoid absent; preopercle ridge smooth, edge serrated at angle or smooth; prootic narrowed along internal orbit rim, not quit excluded by pterosphenoid and parasphenoid; anterior pair of uroneurals absent, posterior pair of uroneurals reduced or absent; two epurals; hypurals 1–2 fused, 3–4 fused, separate from urostyle; hypural 5 free; parhypural free; free medial extrascapular; neuromasts reduced to a simple cross-hatch on head, linear and vertical rows on body, present on caudal fin; cephalic pore system simple without canal flutes; lateral-line scales 23–43, first few scales pored, followed by notched scales, a lower lateral line of notched scales begins on abdomen; cycloid scales on body.

Distribution. Members of the Pseudamiinae are found along the continental coasts and islands of the Red Sea, Persian Gulf, Indian Ocean, Western Pacific out to Japan, Palmyra, Tahiti, Austral Islands and Australia. They can be found in shallow water down to about 64 meters.

Remarks. This subfamily contains one genus, 7 species: Pseudamia amblyuroptera ( Bleeker 1856c) , P. gelatinosa Smith 1956 , P. hayashii Randall, Lachner & Fraser 1985 , P. nigra Allen 1992 , P. rubra Randall, Lachner & Fraser 1985 , P. tarri Randall, Lachner & Fraser 1985 , P. zonata Randall, Lachner & Fraser 1985 . The molecular analyses support a separate subfamily for species of Pseudamia (see Figs. 2–6 ). Morphological information associated Gymnapogon and Pseudamiops with Pseudamia ( Smith, 1961; Fraser 1972 ; Baldwin & Johnson 1999; Bergman 2004) as belonging in this subfamily. However, Thacker & Roje's (2009), Cowman & Bellwood's (2011), and the present molecular analyses placed Pseudamia outside the rest of the Apogonidae with Gymnapogon nested into the latter large clade ( Pseudamiops was absent from these studies). Corresponding to such phylogenetic relationship, monophyly between Pseudamia and Gymnapogon (H14) was significantly rejected by the AU test based on the present molecular data ( Table 6 ). All Pseudamia have two scaled lateral lines on the body, hypurals 1 and 2 fused, urostylar sheath over hypurals 3 and 4, one reduced pair of uroneurals in anterior position, all possible synapomorphies. The comparative morphological features of Pseudamia are reductive, fusion and loss of bones ( Baldwin & Johnson, 1999). Many of these reductive features are held in common with Gymnapogon , Cercamia Randall & Smith 1988 , Lachneratus Fraser & Struhsaker 1991 and Pseudamiops . The latter four genera are all translucent when alive unlike most (all?) species of Pseudamia . Body shapes of Gymnapogon , Cercamia and Lachneratus include forked caudal fins and the latter two genera have large scales and lack pored lateral-line scales. Pseudamiops was not part of this study and is removed from the Pseudamiinae to the Gymnapogonini based on its body being translucent, having large scales without pored lateral-line scales. The conflict between using molecular and morphological information independently supporting differing hypotheses remains confounding for these five genera.

Living pseudamine fishes have virtually no morphological characters that are considered basal for apogonids as restricted here. All are derived characters from those states present in the Amioidinae and in the basal Apogoninae . Baldwin & Johnson (1999) listed plesiomorphic characters for the expanded pseudamine fishes. They did not include Lachneratus or Cercamia as part of the pseudamine group.

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.