Review of the cardinalfishes of the genus Cercamia (Percomorpha: Apogonidae) of the Red Sea and Indian Ocean with descriptions of three new species.

The representatives of Cercamia from the Indian Ocean including Red Sea are reviewed and three new species are described: Cercamia spio n. sp., formerly known as C. eremia (Allen, 1987), is described from 14 specimens, 1733 mm SL, collected in 1015 meters from northern (Duba) to central (Jeddah) Saudi Arabia and from Jezirat Faraun, Egypt. It also has been photographed from the Gulf of Aqaba (Dahab, Egypt) and El Quseir (Mangrove Bay, Egypt). The new species is distinguished from other Indian Ocean Cercamia in having fewer developed gill rakers on lower limb (usually 11 versus usually 1213) and fewer anal-fin rays (11 versus usually 1213). Another new species, Cercamia laamu, n. sp., is described only from the Maldives and Chagos Archipelago based on five specimens 16.030.5 mm SL. It differs from all Indian Ocean Cercamia in having more greater number of the second dorsal-fin rays (10 versus usually 9), and a translucent body devoid of reddish marks versus small reddish dots and crisscross lines. The third new species, Cercamia mascarene, n. sp., is described from 40 specimens 1936 mm SL, from Rodrigues Island, Mauritius. It differs from Cercamia eremia in having a greater number of developed gill rakers on the lower limb of the first gill arch (usually 13 versus usually 12). Free neuromasts and cephalic pores are illustrated for Cercamia mascarene and free neuromasts on the body and caudal fin are illustrated for Japanese specimens of C. cf. eremia. New diagnoses are provided for Cercamia cladara, the type species of the genus, and C. eremia. A map of collection locations for species of Cercamia is presented to show the breath of known occurrences in the Red Sea and Indian Ocean. A new morphologic diagnosis is provided for Cercamia. A phylogenetic analysis of the barcoding portion of the mitochondrial COI gene, including all available sequences from members of the genus Cercamia, displays a much higher species diversity than expected, with high levels of divergence among recognized and undescribed species. A key to the described Indian Ocean species is provided.

Status of the cardinalfish genus Apogonoides and its type species Apogonoides macassariensis (Percomorpha: Apogonidae).

Apogonoides Bleeker 1849 is identified as the senior synonym of Cercamia Randall Smith 1988. We present evidence that Cercamia has become well established in the literature. Reversal of priority is proposed for the genus. The type species, macassariensis, is identified with Cercamia eremia (Allen 1987) based on geographic evidence. Reversal of priority is proposed for the species.

A new species of cardinalfish (Gymnapogon, Gymnapogonini, Apogonidae, Percomorpha) from the Philippines.

The new species is consistent with Gymnapogon, a distinct genus having one to three spines on the preopercle edge, fused hypurals (parhypural+1+2 and 3+4+terminal central), a free fifth hypural, two epurals, no supraneurals, scaleless head and body, a single rod-like postcleithrum and complex lines of free neuromasts on the head, body and caudal fin. The new species is distinguished by having a combination of a rounded caudal with 15 branched principal caudal-fin rays, 2 unbranched principal caudal-fin rays, 10 soft dorsal rays with the anterior two rays unbranched, 9 or 10 anal rays with the first ray unbranched and 14 pectoral rays the lower three and upper two unbranched. Nominal species Gymnapogon          annona, G. foraminosus, G. japonicus, G. urospilotus and G. vanderbilti have 9 or 10 soft dorsal, 9 or 10 anal rays and 12 to 14 pectoral rays, the lower two and upper two unbranched. Gymnapogon africanus, G. melanogaster and G. philippinus have 9 soft dorsal rays, the first ray unbranched and 8 soft anal rays, the first ray branched. Gymnapogon japonicus has a rounded caudal fin with 13, 14 or 15 branched principal caudal rays. Gymnapogon africanus, G. annona, G. melanogaster, G. philippinus, G. urospilotus and G. vanderbilti have a forked caudal fin with 13 branched principal caudal rays and 2 unbranched upper and lower principal caudal rays. A single preopercle spine distinguishes the new species from the single bifid-like preopercle spine shared by Gymnapogon annona, G. melanogaster and undescribed forms. The wide-spread B-marked species complex made up of Gymnapogon urospilotus, G. vanderbilti and undescribed forms have at least one small upper preopercle spine in addition to the larger single spine near the angle of the preopercle.

A new genus and species of cardinalfish (Percomorpha, Apogonidae, Sphaeramiini) from the coastal waters of Vietnam: luminescent or not?

Xeniamia atrithorax is a diminutive new genus and new species. The following combination of melanophore patterns is unique among known apogonids: a large cluster of melanophores in the skin anterior to the insertion of the pelvic-fin base, then extending forward along the sides of the isthmus; posterior portion of the oral chamber is black with melanophores extending below the gill apparatus forward along the base of the oral chamber thence along the basibranchials ending posterior to and below the level of the tongue; a line of large melanophores extends along the inner side of the ceratohyals; black to blackish stomach; and the anterior portion of the intestine is mostly pale with few black spots becoming more densely spotted with melanophores past the first bend and black from the second intestinal turn to the anus. The peritoneum is silvery with scattered melanophores that are more densely distributed along the lower portion of the abdominal cavity. There are two pale, large pyloric caeca at the connection between the stomach and intestine. The combination of eight first-dorsal spines, two supernumerary dorsal spines, one supraneural, lacking an ossified basisphenoid, two epurals and fused hypurals 1+2, preopercle with a smooth ridge and edges, fourth dorsal spine longer than third distinguish this cardinalfish from all other genera. This new genus may be related to Jaydia in the Sphaeramiini rather than with the Ostorhinchini or Siphamiini. Possible luminescent activity is inferred from anatomy and unique melanization patterns versus suppression of light from luminescent prey in the alimentary canal. A table lists suspected and known luminescent apogonids.

Foa yamba, a new species of cardinalfish (Percomorpha: Apogonidae: Apogonichthyini) from the tidal region of the Clarence River, Australia and redescriptions of the West Pacific Foa longimana and Foa hyalina.

Foa yamba, a new species is described from an eastern Australian tidal estuary. This species differs from other West Pacific Foa by having many small spots on the body. Foa longimana, known only from the holotype, is presently an unidentifiable larval stage from Indonesia and is redescribed, but not allocated to a different genus. Foa hyalina, a West Pacific species is reviewed and its known distribution expanded. It is distinguished by five reddish or brownish-red bars on the body from nape to the level of middle portion of the second dorsal fin and lacks markings on caudal fin. 

A new genus of cardinalfish from tropical Australia and southern New Guinea (Percomorpha: Apogonidae).

Ozichthys is described as a new monotypic genus, type species Apogon albimaculosus. This species is restricted to tropical Australia and southern New Guinea. The combination of first dorsal fin with VIII visible spines, fused first and second hypurals, epineural on ninth rib, ossified lower preopercle edge, serrated edge of the preopercle, more than three flute canals on post orbit, 23-24 pored lateral- line scales, elongate last lateral-line scale, and sixteen rows of free neuromasts on principal caudal fin-rays will identify this genus from other brownish ground-colored apogonids. This genus is placed in the clade with Apogonichthys, Foa, Fowleria, and Neamia based on color patterns present on the head, body and vertical fins and in having a single pore above and below the raised median canal on each pored lateral-line scale. 

Revision of the systematics of the cardinalfishes (Percomorpha: Apogonidae) based on molecular analyses and comparative reevaluation of morphological characters.

Molecular analyses were conducted based on 120 of the estimated 358 species of the family Apogonidae with 33 of 40 genera and subgenera, using three gobioids and one kurtid as collective outgroups. Species of Amioides, Apogon, Apogonichthyoides, Apogonichthys, Archamia, Astrapogon, Brephamia, Cercamia, Cheilodipterus, Fibramia n. gen., Foa, Fowleria, Glossamia, Gymnapogon, Jaydia, Lachneratus, Nectamia, Ostorhinchus, Paroncheilus, Phaeoptyx, Pristiapogon, Pristicon, Pseudamia, Pterapogon, Rhabdamia, Siphamia, Sphaeramia, Taeniamia, Verulux, Vincentia, Yarica, Zapogon and Zoramia were present in the molecular analyses; species of Bentuviaichthys, Holapogon, Lepidamia, Neamia, Paxton, Pseudamiops and Quinca were absent from the analyses. Maximum-likelihood (ML), Bayesian (BA), and Maximum parsimony (MP) analyses based on two mitochondrial (12S rRNA-tRNAVal-16S rRNA, ca. 1500 bp; COI, ca. 1500 bp) and two nuclear DNA (RAG1, ca. 1300 bp; ENC1, ca. 800 bp) fragments reproduced two basal clades within the monophyletic family: one including a single species, Amioides polyacanthus, and the other comprising species of Pseudamia. All the other apogonid species formed a large well-established monophyletic group, in which almost identical 12 major clades were reproduced, with phylogenetic positions of four species (Glossamia aprion, Ostorhinchus margaritophorus, Pterapogon kauderni, and Vincentia novaehollandiae) left unsettled. Apogon sensu lato and recent Ostorhinchus (excepting O. margaritophorus) were divided into six and three major clades, respectively. Each of the recognized clades in the family was then evaluated for morphological characters to identify synapomorphies. Based on the results of the molecular analyses and the reevaluation of morphological characters, four subfamilies were proposed within the family: Apogoninae (including most of the species in the family), Amioidinae new subfamily (including Amioides, and based on morphology, Holapogon), Paxtoninae new subfamily (including Paxton, based only on morphology) and Pseudamiinae (including Pseudamia). Within the largest subfamily Apogoninae, twelve new tribes were proposed based on the 12 molecular clades and associated morphology: Apogonichthyini, Apogonini (mainly including species of Apogon sensu stricto), Archamiini, Cheilodipterini, Gymnapogonini, Ostorhinchini (including striped species of recent Ostorhinchus), Pristiapogonini, Rhabdamiini, Sphaeramiini (mainly including barred species of traditional Ostorhinchus, such as Apogonichthyoides, Jaydia and Nectamia), Siphamiini, Veruluxini, and Zoramiini. Two additional tribes are proposed based only on morphology: Glossamiini and Lepidamiini. For each of the 14 tribes, morphological characters were described. One new genus, Fibramia, type species Apogon thermalis, recently in Ostorhinchus, was described supported by morphology and molecular trees. A key to all genera is provided and all valid and uncertain status species are allocated to tribes and genera.

A new genus of cardinalfish (Apogonidae: Percomorpha), redescription of Archamia and resemblances and relationships with Kurtus (Kurtidae: Percomorpha).

Archamia is restricted to a single species, A. bleekeri. A recently described genus, Kurtamia, a reference to a suggested relationship with the enigmatic Kurtus, is the junior synonym of Archamia. Kurtamia bykhovskyi is a junior synonym of A. bleekeri. Archamia is redescribed using osteological, color pattern, pore and free neuromast patterns supplementing those characters used in other publications noting clear differences between A. bleekeri and all other species formerly in that genus. The new genus Taeniamia, type species Archamia leai contains the remaining species. Osteology, color patterns and lateralis characters are reviewed for Taeniamia leai and other species. Species placed in Taeniamia have two broadly different color patterns: 1) yellow, orange, red or dark bars with or without a dark basicaudal spot, and 2) yellow or dark midline stripe with another stripe above the lateral line, lacking bars. These color patterns suggest two lineages exist within Taeniamia. New species combinations are: Taeniamia ateania, T. biguttata, T. bilineata, T. buruensis, T. dispilus, T. flavofasciata, T. fucata, T. kagoshimana, T. leai, T. lineolata, T. macroptera, T. melasma, T. mozambiquensis, T. pallida, and T. zosterophora. Archamia and Taeniamia are sister genera. A diagnosis is provided for the Apogonidae: one or two anal spines, first spine small, supernumerary in position, second spine or first anal fin-ray (only Paxton) in serial association with first distal and proximal-middle radials; first segmented anal ray branched; males mouth brood fertilized eggs; swim bladder simple without anterior or posterior modifications, a dorsal oval and ventral gas glands; free neuromasts on head, body and caudal fin. Characters of Holapogon were used to help identify common plesiomorphic characters for the Apogonidae, elsewhere among percoids using the Centropomidae and information for basal Percomorpha. A table of basal characters and derived changes is provided for the Apogonidae. Characters for two species Kurtus indicus and K. gulliveri are described and examined in a search for morphological synapomorphies with Archamia, Taeniamia, Holapogon and other apogonids. A diagnosis is provided for the Kurtidae: highly modified ribs, anterior dorsal spines individually fused with all radials of the pterygiophore complex, medially fused pterosphenoids, gill rakers on second branchial arch, tooth plates between each gill raker, serrated curved extension of the male's supraoccipital crest, tiny cycloid scales on head and body, very short pored lateral-line scales and free neuromast patterns on the head and body. The second epibranchial articulating with third pharyngobranchial and radial ridges simple or bifid filaments around the micropyle of the egg related to egg ball organization are supported as possible non-exclusive synapomorphic characters for kurtids and apogonids. Parental care of eggs has not been demonstrated for Kurtus indicus, an estuarine/coastal marine species. Kurtids share foramina in the lateral lower part of each caudal vertebra with carangine species, some ephippids, some leiognathids, some priacanthids and scatophagids and share tiny cycloid scales with carangoids: possible synapomorphies or independently derived features.