Sphaeramiini new name Fraser, 2014

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 : 186

Tribe Sphaeramiini new name Fraser & Mabuchi

Type genus Sphaeramia Fowler & Bean 1930

Diagnosis. Members of the Apogoninae : dorsal fin VII+I,9–10, VII(I)+I,9–14 or VIII+I,9–10 with eighth spine reduced to a tiny visible spine or a nubbin hidden under skin supported by a free sixth distal radial; anal fin II,8–13; head and body with ctenoid scales; pored lateral-line scales 23–27; preopercle ridge smooth, edges serrate; 2–3 supraneurals; supramaxilla absent; basisphenoid present; one pair of uroneurals; three epurals; five free hypurals or 1+2 fused and 3+4 fused with 3+4 fused to terminal centrum; parhypural separate; two autogenous haemal spines; two supernumerary dorsal spines; caudal fin forked, emarginate or truncate.

Other characteristics. first segmented fin-ray in second dorsal-fin branched, first anal-fin ray branched and segmented; ctenoid scales on predorsal, cheek, breast, two pelvic scales, and body; ctenoid scales on opercle and onto base of caudal fin; 1–3 predorsal scales cycloid or ctenoid; pored lateral-line scales with multiple pores; pectoral-fin rays 13–17; 9+8 segmented principal caudal rays, 15 branched, upper and lower unbranched; developed gill rakers 7–38; unbranched procurrent rays, longest segmented or spinous; third or fourth dorsal spine longest; 10+14 or 10+15 vertebrae; 8 ribs; 8–9 epipleurals; teeth in one or multiple rows on premaxilla, dentary, vomer, palatine, all villiform; six infraorbitals, bony shelf on third infraorbital; stomach and intestine pale or black with pale peritoneum; low crest on PU2.

Distribution. Species of Apogonichthyoides are found throughout the Indo-West Pacific from East Africa, Red Sea, India, some island of the Indian Ocean, throughout Indonesia, Philippines to Japan, New Guinea, Australia and the Solomon Islands. Some species have restricted distribution and some are rare in collections. Species of Jaydia Smith 1961 have a more continental distribution, mostly caught in trawls, and are widespread from Africa to Japan, New Guinea, Australia and larger islands in the Coral Sea. Representatives of Nectamia are found on reef habitats from East Africa, Red Sea, islands in the Indian Ocean, throughout the West Pacific to Japan, onto the Pacific Plate out to the Line Islands and French Polynesia. Sphaeramia , represented by two species, one found in mangrove habitats and one on coral reefs, also have wide distribution in the Indo-West Pacific. Quinca , represented by a single species, is known from northwestern coast of Australia.

Remarks. This tribe contains six genera, Apogonichthyoides , Jaydia , Nectamia , Pterapogon Koumans 1933 , Quinca and Sphaeramia , almost corresponding to the clade IV in the molecular trees ( Figs. 2–6 , Table 4 ). First three of the six genera were formerly classified under Ostorhinchus , corresponding to "barred group" of Ostorhinchus ( Ostorhinchus II ) in Mabuchi et al. (2006). Two of the remaining three genera ( Pterapogon and Sphaeramia ) have distinct bar(s) on body. In addition to the six genera, corresponding molecular clade included three species of Ostorhinchus -2 ( O. hoevenii Bleeker 1854b , O. ishigakiensis Ida & Moyer 1974 , and O. rueppellii Günther 1859 ). But there are no morphological characters that support this association. We defer taxonomical action on this possible relationship for the present. This clade further included another species of the former Ostorhinchus ( Ostorhinchus -1). For this species, the latest authors ( Allen & Erdmann 2012) provisionally used Apogonichthyoides as its genus name, and we followed it here, but such application of the name made this genus paraphyletic in our molecular trees. Based on its morphological features, we tentatively included it [ Apogonichthyoides (?) melas ] in this tribe, but further study including more Sphaeramiini species will be needed to clarify its taxonomic status. In the present molecular analyses, phylogenetic position of Pterapogon remained unsettled ( Table 5 ). It was sister to the clade VII ( Siphamiini ) in ML tree ( Fig. 2 ), sister to Ostorhinchus margaritophorus in MP tree-A ( Fig. 4 ), and nested within the clade XII ( Ostorhinchini ) in the MP tree-B ( Fig. 5 ). It formed a polytomy with O. margaritophorus , clade VII, clade VIII, and a large clade including clades IX–XII in BA tree ( Fig. 3 ). Pterapogon has, however, a synapomorphy (spinous procurrent caudal rays) uniquely held with Sphaeramia as well as similarities in color patterns and body shape, but differs with fused hypurals 1+2 and 3+4. Based on these morphological features, we tentatively included this species in the tribe Sphaeramiini . Monophyly of them was not rejected by AU test (H02 in Table 6 ), although its inclusion within Ostorhinchini was also not rejected (H03 in Table 6 ). We had no good DNA extracts from tissue of Quinca . This monotypic genus has morphological characteristics and color pattern suggestive of a relationship with Apogonichthyoides , and was provisionally place in the Sphaeramiini .

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.