Taeniamia, Fraser, 2013

Fraser, Thomas H., 2013, A new genus of cardinalfish (Apogonidae: Percomorpha), redescription of Archamia and resemblances and relationships with Kurtus (Kurtidae: Percomorpha), Zootaxa 3714 (1), pp. 1-63 : 18-30

Taeniamia new genus

Type species Archamia leai Waite, 1916

This apogonine genus has at least 14 species.

Figures 1–5 , 6–8 , 10 , 12 , 14–19 , Table 1

Material examined: Taeniamia ateania : Archamia ateania Holotype BPBM 37600 , (1, 39.5), Siberut Island , Sarabua Bay, Sumatra, Indonesia, 6 m. Paratypes CAS 98472, (1, 41.4) & BMNH 1998.1.6.1, (1, 40.3), Mentawai

Is., Sumatra, Indonesia. USNM 347598, (1, 44.8), Ko Surin, Andaman Sea, Thailand, digital x-ray. Taeniamia biguttata : Archamia biguttata Lectotype USNM 56156, (1, 60), Bacon, Philippines, digital from film x-ray. Paralectotypes USNM 360647, (1, 60.6), cleared and stained. USNM 360647 (4, 56.6–63.7), same data as lectotype, digital and film x-ray. USNM 112158, (5, 49.0–57.0), vicinity of Surigao Strait, Leyte, San Roque, Philippines, digital from film x-ray. USNM 126370, (6, 27–59), Bacon, San Fabian, Philippines, digital of film x-ray. Taeniamia bilineata : Archamia bilineata Holotype BPBM 21514, (1, 30.9), El Hamira, Sinai, Egypt. Paratypes USNM 331828, (1, 29.6), same data as holotype, digital from film x-rays. Taeniamia buruensis : Apogon buruënsis Holotype RMNH 5562, (1, 46.7), Kajeli, Buru I., Moluccas Is., Indonesia. USNM 112111, (10, 50–66), Illana Bay, S of Mindanao River, Philippines, digital from film x-ray. USNM 112112, (1, 62), Nunucan River, Mindanao, Philippines, cleared and stained. Taeniamia dispilus : Archamia dispilus Holotype, USMN 112041, (1, 57.6), Soo Wan Bay, Taiwan, digital from film x-ray. Paratypes USNM 112077, (5, 53–68), same data as holotype, digital of film xray, one 53, cleared and stained. USMN 112078, (1, 57), Kwa Siang Bay, Taiwan, digital from film x-ray. Taeniamia flavofasciata : Archamia flavofasciata Holotype SAIAB 3395, (1, 48.8), Durban, South Africa, digital of film x-ray. Paratypes: BMNH 1936.8.7.2–7, (2, 55.4–59.7), Mombasa, Kenya. BMNH 1867.3.9.558, (1, 61.3), Zanzibar, Tanzania. SAIAB 62641, (2, 47–49), same data as holotype, digital from film x-ray. USNM 307710, (2, 55.9–56.8), Nosy Bé, Madagascar, x-ray. Taeniamia fucata : Apogon fucatus Holotype BMNH 1860.3.19.353, (1, dried skin, one side), Pinang, Malaysia. Apogon macropteroides Holotype RMNH 5608, (1, 58.5), Lepar I., Indonesia. Archamia irida Paratype BPBM 18184, (1, 44), Gulg of Suez, Egypt. Paratype, USNM 331829, (1, 33.5), Gulf of Suez, Egypt, digital from film x-ray. SAIAB 3400, (1, 54), Mahe I., Seychelles, digital of film x-ray. SAIAB 3402, (1, 52), Aldabra, digital of film x-ray. SAIAB 3404, (1, 63), 3408, (1, 62), Zanzibar, digital from film x-ray. SAIAB 3407, (3, 38–61), Malindi, Kenya, digital of film x-ray. SAIAB 3411, (1, 70), Shimoni, Kenya, digital from film x-ray. Mozambique (all digital from film x-rays): SAIAB 3401, (1, 60), 3403, (4, 52–59), Mozambique I. SAIAB 3405, (4, 42–69), SAIAB 3409 (2, 63–66), 3415, (1, 50), 3417, (9, 53–64), Pinda I. SAIAB 3406, (2, 45–45), Kifuk I. SAIAB 3410, (3, 61–63), Mozambique I. SAIAB 3412, (1, 61), 3414, (2, 54–54), Inhaca I. SAIAB 3413, (4, 47–56), Wamizi I. SAIAB 3416, (6, 54–65), Rolas I. UF 173573, (1, 57), Sri Lanka. USNM 356401 (3 of 822), Pulau Seribu, Indonesia, 5°51'S 106°34'E, digital x-ray. USNM 52203, (1, 55), Apia, Samoa, cleared and stained. USNM 111967, (5 of 11, 54–57), Samoa, Apia, digital from film x-ray. 149405, (5, 59–62), Tutu Bay, Jolo I., Philippines, digital from film x-ray. Taeniamia leai : Archamia leai , Syntypes SAMA F 308, (4, 58.4–67.9), Norfolk Island, partially dried out, 68 mm, digital from film x-ray. USNM 339371, (2, 54–56), One Tree I., Queensland, Australia, digital from film x-ray. USNM 356269, (3, 53–65), One Tree I., Queensland, Australia. USNM 356272, (1, 53), One Tree I., Queensland, Australia, digital x-ray. USNM 306823 (1, 55.2), No data, cleared and stained. Taeniamia lineolata: USNM 166969, (3 of 15, 24–48), Ghardaqa, Red Sea, Egypt, digital x-ray. USNM 356412, (3 of 10, 32–52), Ghardaqa, Red Sea, Egypt, 27°18'50"N 33°47'35"E, digital x-ray. Taeniamia macroptera : Apogon macropterus Paralectotype, RMNH 51, (1, 66.2), Java, Indonesia. Apogon argenteus Holotype MNHN 8755, (1, 47.4), Vanikoro, Solomon Is., Papua New Guinea. USNM 171228, (1, 53), Puerto Princessa, Palawan I., Philippines, cleared and stained. USNM 171230, (3 of 9, 56–58), Ragay Gulf, Alibijaban I., Luzon, Philippines, digital from film x-ray. USNM 171238, (4, 52–54), Port Ciego, N Balabac Strait, Philippines, digital from film x-ray. USNM 171239, (3 of 9, 37–50), vicinity of Tomahu I., S. Bouro I., Philippines, digital from film x-ray. Taeniamia melasma : Archamia melasma Holotype AMS IB.4473, (1, 59.9), Yirrkala, Northern Territory, Australia, digital from film x-ray. Paratype USNM 173794, (2, 59–60), Yirrkala, Northern Territory, Australia, digital x-ray. Taeniamia mozambiquensis : Archamia mozambiquensis Holotype SAIAB 349, (1, 64.6), Mozambique I., Mozambique, digital from film x-ray. Paratypes: SAIAB 1062, (1, 68.9), Mozambique I., Mozambique, cleared and stained. SAIAB 769, (4, 51–74), digital from film x-ray. SAIAB 770, (2, 71–78) and SAIAB 771, (1, 72), Inhaca I., Mozambique, digital from film x-ray. SAIAB 772, (9, 56–74), digital from film xray. SAIAB 773, (7, 64–73), Mozambique I., Mozambique, digital from film x-ray. BMNH 1864.11.15.7, (2, 38.8– 43.7). Non-type material: USNM 356262, (1, 52), Dar-Es-Salaam fish market, Tanzania, digital x-ray. Taeniamia pallida : Archamia pallida Holotype BPBM 361158 (1, 33.2), Masirah I., Oman, digital from film x-ray. Paratypes BMNH 1994.10.4.2 (1, 28.2), same data as holotype. USNM 331831, (1, 32.0), same data as holotype, digital from film x-ray. Taeniamia zosterophora : Apogon zosterophorus Lectotype RMNH 5600, (1, 41.2), Manado, Sulawesi, Indonesia. USNM 205666 (1, 42), Pulau Gaya, Darvel Bay, Malaysia, cleared and stained. USNM 123386, (2, 31– 42), digital from film x-ray. USNM 171323, (5 of 21, 50–53), Rapu-Rapu I., San Bernardino Strait, E. Luzon, Philippines, digital from film x-ray. USNM 171346, (5, 45–48), digital from film x-ray.

Diagnosis. Hypurals 1–4 free; a slender pair of uroneurals; urostylar sheath over hypurals 3 and 4; ribs without wide flanges; soft anal-rays 11–19; basicaudal dark spot variable in size and intensity; body translucent or semitranslucent in life, with stripes or bars on head and/or body.

Description. Body relatively deep and compressed except T. bilineata ( Gon & Randall 1995) ; eye large; mouth large and oblique except straight in T. bilineata ; oblique; posterior edge of premaxilla concave, straight in T. bilineata ; eye large; mouth slightly oblique, two dorsal fins ( Fig. 1F–K ).

Dorsal-fin VI–I,9 or VII–I,9; spines slender, none robust, as VI(0)–I or VII(0)–I, first dorsal fin with VII visible spines (only T. leai , Figs. 2B & 3B ) or VI (all other species as in Fig. 3A but without shorter anterior spine); three supraneurals, first anterior to first neural spine with second between neural spines 1 and 2 and third between neural spines 2 and 3, all supraneurals with slightly expanded tips; first proximal-middle radial without procumbent spine, first proximal-middle radial between neural spines 2 and 3, second and third proximal-middle radial between neural spines 3 and 4; one or two supernumerary spines associated with first proximal-middle radial, first supernumerary spine (I) very short, second supernumerary spine (II) less than half length of third spine ( T. leai only, Fig. 2B ), or single supernumerary spine (homologous as supernumerary spine II), less than half length of second spine (homologous with spine III, all other species); middle radials fused to proximal radials 1–12, free for middle radials 13–16; distal radials 1–3 sutured or with processes associated with proximal-middle radial, free for middle radials 4–16, proximal-middle radial 6 with long distal arm, its distal radial on dorsal edge of seventh proximal-middle radial ( Fig. 3E, T . leai) or fused with dorsal edge of seventh proximal-middle radial (like Fig. 3D ) and not bearing remnant spine or nubbin below skin [e.g., T. buruensis ( Bleeker 1856a) , ( T. dispilus Lachner 1951 ), T. fucata , T. macroptera , T. zosterophora ( Bleeker 1856b) ]; last distal radial with split fin-ray supported by bony stay ( Fig. 6A ). See table 1 for axial skeleton relationships between haemal spines and anal fin elements.

Anal-fin spines, single supernumerary spine, second spine serially associated with first proximal-middle radial ( Fig. 4B ); first proximal-middle radial much longer than second both with rod-like tips, in close association with last pair of ribs and first haemal spine; first proximal-middle radial in advance of haemal spine on centrum 11, second proximal-middle radial behind or in front of first haemal spine; distal expansion as anterior flange on first proximal-middle radial; first proximal-middle radial not articulating with centrum 10 ( Figs 4B , 5A ); first proximalmiddle radial with curved anterior flange ( Fig. 4B ); proximal-middle radial formula for first haemal spine and following interhaemal gap as 1/4 (1 of 1 T. ataenia ( Randall & Satapoomin 1999) , 15 T. biguttata (Lachner 1951) , 1 of 6 T. dispilus , 1 of 3 T. flavofasciata Gon & Randall 2003b ), 41 of 55 T. fucata , 2 of 7 T. leai ( Fig. 4B ), 3 of 6 T. lineolata , 6 of 10 T. macroptera ( Fig. 5A ), 3 of 3 T. melasma ( Lachner & Taylor 1960) , 3 of 25 T. mozambiquensis ( Smith 1961) , 6 of 9 T. zosterophora ); 1/5, (1 of 1 T. bilineata , 5 of 7 T. leai [also see Springer and Smith-Vaniz, 2008:32], 1 of 10 T. macroptera , 7 of 25 T. mozambiquensis ); 2/3 (5 of 10 T. buruensis , 5 of 6 T. dispilus , 2 of 3 T. flavofasciata , 14 of 55 T. fucata , 3 of 6 T. lineolata , 3 of 10 T. macroptera ,1 of 25 T. mozambiquensis , 3 of 9 T. zosterophora ); 2/4 [5 of 10 T. buruensis ,14 of 25 T. mozambiquensis , 1 of 1 T, pallida ( Gon & Randall 1995) ]; soft anal fin-rays all branched, range 12–19, usually 12 ( T. buruensis ), 13 ( T. bilineata , T. flavofasciata , T. leai , T. lineolata , T. pallida ), 14 ( T. macroptera , T. mozambiquensis ), 15 ( T. ataenia , T. zosterophora ), 16 ( T. biguttata , T. dispilus , T. fucata ) 17 ( T. dispilus ); usually small proportion of variation either side of common count (except T. biguttata , T. zosterophora ); middle radials fused to proximal radials 1–7, free on 8–13; distal radials all free; last distal radial with split fin-ray supported by bony stay ( Fig. 6E ).

Vertebrae 10+14 ( Fig. 2B , Table 1); open haemal arches on vertebrae 6–10 ( Figs. 4B , 5A & 7B ); arch on sixth vertebra short and narrow, arches on seventh to ninth vertebrae progressively longer ( Fig. 4B & 7A ), tenth arch narrow, without flaring; epineurals articulate with vertebrae 1–2, then with ribs on vertebrae 3–8, absent on last two ribs ( Fig 7B ); ribs on third to tenth vertebrae; first rib slender, rod-like its proximal tip articulating on neural arch of third vertebrae; second rib slender rod-like distally, its proximal tip articulating just above mid-centrum ( T. leai and T. macroptera with slight proximal posterior expansion); third rib with flattened posterior edge for T. leai ( Fig. 7C ), not shown for T. dispilus , T. fucata , T. zosterophora , otherwise rod-like, its proximal tip articulating just below mid-centrum; fourth rib with posterior edge slightly expanded (not illustrated, T. buruensis , T. dispilus , T. furcate , T. zosterophora ) otherwise rod-like, articulating on very short haemal arches; fifth to seventh ribs rod-like, articulating with progressively longer haemal arches (proximal expansion on both edges, T. buruensis , T. dispilus , T. furcate , T. zosterophora , narrow posterior expansion only for T. macroptera ); eighth rib short, flange-like without narrow rod-like distal end, articulating with longest haemal arch; eighth rib always associated with first anal proximal-middle radial, second anal proximal-middle radial shorter than first, not associated with eighth rib if behind first haemal spine or associated with rib if proximal-middle radial anterior to first haemal arch.

Hypural complex ( Figs. 2B & 8B ): five free hypurals; free parhypural with long apophysis; three epurals, first two with expanded proximal ends; first pair of slender uroneurals overlapping proximal half of fifth hypural and distal half of urostyle ( T. leai ) or shorter just reaching past proximal end of fifth hypural, second pair of uroneurals absent; second and third preural centrum with ( T. dispilus , T. furcate , T. macroptera , T. zosterophora ), or ( T. leai ) without autogenous haemal spines; low neural crest on preural 2; neural and haemal spines on third preural centrum thicker and longer than spines on fourth preural centrum; dorsal and ventral procurrent ray 8–9 and 7–8 respectively, not spinous, articulating with cartilages, haemal spine of second preural centrum and epurals, no procurrent spur; 9+8 principal caudal rays, articulating with third preural centrum haemal spine to fifth hypural, 8+7 branched; caudal fin forked, some small scales on base of rays.

A single pelvic spine and five branched rays, inner edge of fin with short attachment to body; spine with a rodlike projection on inner sides, not meeting through pelvic foramen.

Pectoral girdle (not shown): Posttemporal with small serrae, arms about same length articulating with neurocranium, extrascapular with four foramina, two in each anterior arm; supracleithrum articulating with posttemporal, overlapping upper portion of cleithrum; coracoid and scapula separate, closely associated with cleithrum; scapula with large central foramen, smaller foramina near posterior upper corner; strut of coracoid meets inner side of cleithrum about 20% up from ventral tip of cleithrum; large upper postcleithrum mostly covered by expanded cleithrum; lower postcleithrum barely articulating with upper postcleithrum; four radials with one articulating with coracoid and three with scapula; pectoral fin-rays with 1–2 upper unbranched ray and 1–3 lower unbranched rays, usually 13 ( T. bilineata , T. zosterophora ), 14 ( T. ataenia , T. biguttata , T. buruensis , T. dispilus , T. flavofasciata , T. fucata , t. lineolata , T. macroptera , T. melasma , T. pallida ) or 15 ( T. leai 2 upper and 2 lower unbranched); usually small proportion of variation in counts on either side of common count.

Ventral gill arches (not shown): proximal cartilaginous end of first basibranchial articulates with anterior end of second basibranchial and sandwiched between paired anterior ends of first hypobranchials cartilaginous end of second basibranchial articulates with anterior end of third basibranchial dorsal to cartilaginous tips of paired anterior ends of second hypobranchials; third basibranchial long, partly sandwiched between the second hypobranchial, posterior end ventral to the small squarish third hypobranchial with long narrow cartilage extending toward the paired fifth ceratobranchials.

Dorsal gill arches ( Fig. 10B ): first (suspensory) pharyngobranchial ossified with cartilage tips, connection to cartilage tip of first epibranchial; interarcual cartilage from unicinate process of first epibranchial dorso-lateral to end of second pharyngobranchial, second pharyngobranchial proximal cartilage tip; distal end of second epibranchial rising and curving over proximal end of second pharyngobranchial to articulate with medial cartilage on edge of third pharyngobranchial; third pharyngobranchial large, proximal cartilage end articulating with distal ends of third and fourth epibranchials; third epibranchial partially Y shaped with dorsal flange curved slightly over fourth epibranchial; fourth epibranchial with low flange; fourth pharyngobranchial absent, a small tooth plate supported dorsally by the fourth epibranchial.

Gill arch dentition: gill rakers on outer side of hypobranchial, ceratobranchial and epibranchial of first arch, rudiments on inner edges; first pharyngobranchial without tooth plate or rudiments; second arch with rudiments and pseudo-rakers on outer side of hypobranchial, ceratobranchial and epibranchial, rudiments on inner edges of hypobranchial, ceratobranchial and epibranchial, second pharyngobranchial with long, narrow tooth plate; tooth plates and rudiments on both sides of hypobranchial, ceratobranchial and epibranchial of third arch, large tooth plate covering most of third pharyngobranchial; small tooth plate on outer side of fourth ceratobranchial, roundish tooth plate near distal end of fourth epibranchial, fourth pharyngobranchial absent; fifth ceratobranchial with large tooth plate over most of its surface, inner posterior third with row of larger teeth than on rest of tooth plate. First gill arch rudiments and raker all species combined: upper rudiments 1–4, upper rakers 2–4, lower rakers 13–18 lower rudiments 0–3, all rakers 19–26, rakers and rudiments 19–27.

Hyoid series: Seven branchiostegals, anterior three on ventral edge of anterior ceratohyal, two on face of anterior ceratohyal, two on face of posterior ceratohyal; anterior ceratohyal deeply notched, foramen absent, with loss of dorsal portion of bone ( Fraser, 1972, Pl. 17B; similar to Fig. 11A ); anterior ceratohyal and posterior ceratohyal with smooth adjoining medial edges; dorsal hypohyal ossified with cartilaginous strip ventrally connecting with ossified ventral hypohyal; basihyal toothless, distal end cartilaginous, proximal end dorsal to dorsal hypohyals with short ligament to upper edge of cartilaginous end of first basibranchial; urohyal wide, with anterior process.

Ventral edge of preopercle serrate or few serrae near angle for T. bilineata , serrate on posterior edge, preopercular ridge smooth, with produced obtuse to acute tip at angle; edge of posttemporal serrate (smooth for T. bilineata ); interopercle, subopercle and opercle smooth.

Six infraorbitals, shelf on third extending along fourth infraorbital but not attached ( T. leai , Fig. 12B ), third with wide shelf and ligaments attached fore and aft at medial corners ( T. buruensis , T. dispilus , T. fucata , T. macroptera , T. zosterophora ) two foramina in first infraorbital, single foramen in each of remaining infraorbitals ( T. leai , T. macroptera ); two foramina in first infraorbital, one in second, two in third ( T. furcate , then single foramen in each remaining; or one foramen in first infraorbital and two in second infraorbital then single foramen in each remaining infraorbital ( T. buruensis ); distal edges smooth.

Neurocranium without oblique crests from distal edge of sphenotic to midline of frontals ( Fig. 14A, T . leai), or with crest from distal edge of sphenotic to midline of frontals ( Fig. 15A, T . buruensis, T. dispilus , T. fucata , T. macroptera , T. zosterophora ); low, short supraoccipital crest ( Fig. 14B T . leai), large supraoccipital crest extending forward over eyes ( Fig. 15B, T . buruensis, T. dispilus , T. fucata , T. macroptera , T. zosterophora ); flanges along midline of frontals aft of mesethmoid; basisphenoid thin ( T. leai ) or well developed ( T. buruensis , T. dispilus , T. fucata , T. macroptera , T. zosterophora ).

Eyes with pair of ossified scleral ossicles ( T. buruensis , T. dispilus , T. fucata , T. macroptera , T. zosterophora ).

No teeth on basihyal, ectopterygoid or endopterygoid; large oval rostral cartilage behind symphysis of premaxilla; maxillary excluded from border of mouth; supramaxilla absent.

Teeth patterns by species: T. leai – teeth on premaxilla villiform with one row; teeth on dentary with slightly enlarge tooth at symphysis, two rows villiform teeth becoming one row on side; 2–3 teeth on each side of vomer; palatine smooth, teeth visible inside bone; no teeth on ectopterygoid. T. buruensis – teeth on maxilla villiform with two to four rows, few near symphysis slightly larger than rest, no taper to fewer rows posteriorly; teeth on dentary villiform band, slightly expanded near symphysis with few inner teeth slightly larger, abruptly becoming single row about half way back on sides. T. dispilus – teeth on premaxilla villiform with one row of slightly enlarged conical recurved teeth near symphysis grading to row of slightly enlarged conical recurved teeth, becoming 2–4 rows of villiform teeth medially tapering to 1–2 rows posteriorly; teeth on dentary villiform with single row of slightly enlarged, conical, recurved teeth at symphysis tapering to slightly enlarged single row medially, smaller posteriorly; vomer with 1 or 2 villiform teeth on each side, none in mid-portion; palatines with long single row of villiform teeth. T. fucata – teeth on premaxilla villiform with one row of slightly enlarged conical recurved teeth near symphysis grading to inner row of slightly enlarged conical recurved teeth, becoming band of villiform teeth medially tapering to 1–2 rows posteriorly; teeth on dentary villiform with 2–4 series of conical recurved teeth at symphysis tapering to slightly enlarged single row medially, smaller posteriorly; vomer with short, single row of villiform teeth on each side, none in mid-portion; palatines with long single row of villiform teeth. T. macroptera – teeth on premaxilla villiform, slightly enlarged conical recurved single row near gap at symphysis, becoming small in 1–2 rows on side; teeth on dentary villiform in two rows, outer row becoming smaller, inner row slightly larger becoming single row on side; vomer with 2–3 recurved teeth; palatine with single long row of villiform teeth. T. zosterophora – teeth on premaxilla villiform with two rows of conical recurved teeth near symphysis becoming narrow band of villiform teeth medially tapering to 1–2 rows posteriorly; teeth on dentary villiform 2 rows of conical recurved teeth, smaller posteriorly; vomer with short, single row of villiform teeth on each side, none in mid-portion; palatines with long single row of villiform teeth.

Scales transforming ctenoid ( Roberts, 1993, App 1), deciduous, often missing in preserved material; pored lateral-line scales ctenoid from posttemporal to base of caudal fin; fourth lateral-line scale ( Fig. 16A ) with large dorsal pore, smaller pore on posterior edge with three large foramina, one small foramen above central canal, three large pores, one inverted dorsally with two large foramina below central canal; predorsal scales 4 ( T. leai ), 1–2 ( T. buruensis ) usually 6 (remaining species); ctenoid scales on cheek, nape, breast, body, two pelvic scales, pelvic axillary scale small or absent, no scales beyond base of pectoral fin, ctenoid scales on base of pectoral fin; caudal fin forked, some small scale on base of rays; pored lateral-line scales 24; row above lateral line 24; transverse row above lateral line 2; transverse row below lateral line 6; circumpeduncular scales 5+2+5 = 12; no scaly sheath at base of anal fin, small scales present between larger ones along fin bases ( T. bilineata , T. leai , T. macroptera , T. pallida ), scaly sheath of small scales at base of anal fin ( T. ataenia , T. biguttata , T. flavofasciata T. fucata , T. lineolata , T. mozambiquensis , T. zosterophora ).

Anterior naris round with short tube, posterior naris oval, close to edge of orbit; naris with underlying simple ossification broadening distally from narrow trough.

Pseudobranchiae present, exposed.

One pyloric caeca.

Cephalic pores patterns incomplete ( Figs. 17A , 18A & 19A ): Snout to nape: anterior supraorbital pore back off edge of snout; series of small pores on central interorbit on sides of area without intervening free neuromasts; two interorbital pores along rim of eye; few pores visible on supratemporal and nape regions. Snout to opercle: lachrymal with an anterior slit-like pore below and behind the anterior naris, two large slit-like pores on lower edge, several small pores under the eye, several small pores along ventral edge above maxilla. Dentary to articular: dentary with an anterior medial pore and ventral medial pore; T. fucata – very numerous micropores similar to T. leai on all parts of head.

Cephalic free neuromasts incomplete ( Fig. 17A ): Dorsal aspect, two irregular rows from preorbit to interorbit with short vertical rows on each side; nape poorly represented; lateral aspect, a long single row on infraorbitals reaching cheek, various short vertical and horizontal rows from around naris, on lachrymal, infraorbitals and onto cheek; ventral aspect multiple longitudinal rows; T. fucata – free neuromast also similar with two longer lateral rows on anterior head, outer one shorter and broadly disjunctive with short line over orbit and another line from upper anterior opercle area to dorsoposterior eye.

Swim bladder with anterodorsal oval; no anterior or posterior modifications associated with neurocranium or first anal proximal-middle radial.

Color patterns: Combinations of following color patterns (see Kuiter & Kozawa, 2001; Gon & Randall, 2003a, b; Allen & Erdmann, 2012): 1) straight, broad bars (1–5) to narrow, straight or slightly curved bars (7–23) on body, bars on opercle; basicaudal spot small to large and diffuse; snout with mark, may extend though eye onto side of head; cheek mark to preopercle; spot/short bar-like mark on body posterior to posttemporal, or 2) combinations of following patterns: one to two stripes, stripe-like series of midline markings; basicaudal spot smaller than eye, usually smaller than pupil; snout with mark, may extend though eye onto side of head.

Stomach and intestine blackish; peritoneum silvery with melanophores.

Etymology. A combination of the feminine Latin noun taenia meaning ribbon or band and the feminine Greek noun Amia meaning a fish, often used with cardinalfishes genera, here as Taeniamia referring to the vertical wide bars and near vertical to curved narrow bars as lines on most of the species in this genus.

Remarks: Some comments about the status of a few species in Taeniamia are presented first then by a discussion of generic characters.

Taeniamia bilineata has some unique characters such as its striped pattern, slender body (likely without frontal crests), mouth and premaxilla shape which suggest a more distant relationship to the other striped species or may belong elsewhere.

Archamia irida Gon & Randall 1995 , a species based on small specimens, was grouped with Archamia bleekeri by Gon and Randall (2003b) based on color pattern, however, they synonymized irida with fucata ( Gon & Randall, 2003a) . Hypural characters seen in radiographs of the paratype support the synonymy.

Taeniamia kagoshimana (Döderlein in Steindachner & Döderlein 1883b) needs additional study. It has the hall marks of a valid species with T. dispilus as a synonym, rather than a synonym of T. fucata . Important characters are: differences in color pattern (see Kuiter & Kozawa, 2001 as Archamia dispilus , p. 101), dentition, gill raker counts (see Gon & Randall, 2003b, table 8, Taiwan to Japan) and flanges on the sixth and seventh ribs. Yoshida et al. (2010, fig. 28) opined that the Japanese species of Archamia they treated was valid but decided to follow Gon & Randall (2003b) and use fucata instead of authors they cited that identified this species as Archamia dispilus . Lachner (1951) described Archamia dispilus from Taiwan and eastern Philippines.

Gill-raker data presented by Gon & Randall (2003b, table 8) also suggest that Taeniamia fucata is a complex of perhaps three species. Distribution of the complex of species/populations may be similar to the following: fucata , ( Maldives, Chagos, Sri Lanka, Indonesia, Philippines, Australia, Palau, Caroline & Marshal Islands, Papua New Guinea, islands out to Tonga), sansibarica ( Pfeffer 1893) (Red Sea, eastern Africa, western Indian Ocean islands) and kagoshimana ( Vietnam, Hong Kong, eastern Philippines, Taiwan, Japan). Bar code molecular evidence would be useful to pursue.

Taeniamia melasma should be recognized as a valid species based on color pattern (see Kuiter & Kozawa, 2003b; Allen & Erdmann, 2012, p. 374) and habitat preference for live Acropora coral rather than a synonym of T. biguttata which is found in caves and under ledges during the day.

Taeniamia has fewer derived character states than Archamia as described above and in the following paragraphs.

In life, the color pattern of Archamia differs from all species of Taeniamia by not having bars or stripes on the head or body, and internally by having yellowish region along the anterior vertebral column and associated peritoneum. A greenish, yellow snout without body bars occurs in juvenile Taeniamia macroptera and juveniles may lack the adult body stripes, appearing similar to Archamia bleekeri . Color patterns are important characteristics ( Mabuchi et al. 2006) for adult cardinalfish expressing overall similarity in patterns among clades supported by molecular evidence.

Fraser (1972) provided osteological illustrations of Taeniamia macroptera . All Taeniamia species have five hypurals, a long pair of slender uroneurals, three epurals, the first epural expanded, all plesiomorphic features at some level for a number of apogonid genera. Taeniamia leai with its two supernumerary dorsal spines, shorter supraoccipital, few body bars and lack of developed oblique frontal crests may represent the living basal Taeniamia . Species in this genus have small proximal flanges appearing on one or both sides of rod-like ribs of the third to seventh vertebrae, a likely synapomorphy among the species. The presence of the proximal flanges on the ribs of among other apogonids has not been well documented. The arrangement and shape of the fifth to seventh dorsal proximal-middle radials may be another synapomorphy shared by Taeniamia and Archamia . Vertical or near vertical first anal proximal-middle radial are shared characters of these two genera and may be another synapomorphy. The number of segmented anal rays (13 or more) was implied by Prokofiev (2006) to be a synapomorphy for Archamia and Kurtamia . However Pterapogon Koumans 1933 , Quinca Mees 1966 and some species of Rhabdamia Weber 1909 have 13 segmented anal rays ( Fraser, 1972) and Taeniamia buruensis has 12. Long anal fins may eventually be shown to be independently derived in Pterapogon , Quinca and Rhabdamia . Supraoccipital crest far forward on the neurocranium, a retrose preopercular spine and a scaly low sheath along the anal-fin base are characters are suggested by Gon & Randall (2003b) as possible synapomorphies. A retrose preopercular spine is not present in Archamia or some of the species of Taeniamia . The supraoccipital crest is high and extends forward in Archamia and Taeniamia with the exception of T. leai . Short, low oblique frontal crests with contiguous half arches are present in Archamia , but absent in T. leai with separated half arches, and well-developed crests in T. buruensis , T. dispilus , T. fucata , T. macroptera and T. zosterophora .

A scaly low sheath on the second dorsal and anal fins is not present in all species ( Gon & Randall 2003b, fig 2).

Gon & Randall 2003b and Gon et al. 2013 report some species with teeth on the tongue. None of the cleared and stained material examined here had teeth on the basihyal. The presence of teeth on the basihyal would be a reversal of a basal condition in apogonids (Table 2).

Those species of Taeniamia with multiple bars, broad or narrow and few or many on the body or present only on the head likely represent a monophyletic group. Archamia ataenia , lacking the body bar of A. zosterophora , is its closest relative ( Allen & Erdmann, 2012, p. 372). None of the other apogonids have bar patterns that can be confused with those found in Taeniamia . The species with stripes on the body likely represent another line as suggested by Gon & Randall (2003b), but this condition is not unique among apogonids. The striped species ( T. bilineata , T. buruensis & T. mozambiquensis ) are maintained in Taeniamia .

Comparing the distribution of cephalic pores of Archamia with Taeniamia leai , there are many more minipores on all aspects of the head of T. leai . The free neuromast pattern is more complex for Archamia in the dorsal aspect, but unavailable for other comparisons. Bergman’s (2004) descriptions and figures of the pores and free neuromasts of Archamia are regarded as incomplete, but present a better coverage of the dorsal and lateral cephalic pores than here.

Fusion in the caudal skeleton, loss of uroneurals, broad expansion of the ribs and a shelf on the third and fourth infraorbital (may or may not occur in other specimens) and long first dorsal spine are apomorphies for Archamia in the clade with Taeniamia . The fusion and loss of elements in the last centrum complex appear elsewhere in unrelated genera (see Fraser, 1972, Fraser & Struhsaker, 1991). Fowleria has shelves on the third, fourth and fifth infraorbital but has no synapomorphic characters to recommend consideration as a close ally of either treated here. All other investigated apogonids either lack shelves or it is only present on the third infraorbital ( Fraser, 1972).

Archamia shares a common apogonid ancestor with Taeniamia based on the long anal-fin base, oblique low frontal crest, vertical first anal pterygiophore, a distal anterior distally-curved flange on the first anal proximalmiddle radial, flanges on both sides of ribs 3–6, all possible synapomorphies with species of Taeniamia . Both genera are derived apogonids based on characters in Table 2. Variation of small or incipient rib flanges among species of Taeniamia is suggestive that this character may be a synapomorphy with Archamia . Pterapogon has narrow flanges (posterior only) on ribs 3–5 and Quinca has narrow flanges (posterior only) on ribs 4–5. Other deep-bodied apogonids are without the lateral frontal crests ( Pterapogon , Sphaeramia Fowler & Bean 1930 , Quinca ) but have well developed oblique crests ( Pterapogon ) and have more numerous soft dorsal- and anal-fin rays ( Pterapogon and Quinca ). Other apogonids with high number of anal-fin rays include Cercamia Randall & Smith 1988 , Lachneratus Fraser & Struhsaker, 1991 , Paxton Baldwin & Johnson, 1999 and Rhabdamia but with more elongate bodies ( Baldwin & Johnson, 1999; Fraser 1972; Fraser & Struhsaker, 1991) and Lachneratus has a high number of soft dorsal rays (12–13).

Figures

FIGURE 1. Archamia bleekeri A–E: A. Durban, South Africa, by D. King. B. El Nido, northern Palawan, Philippines, by G. Allen. C. RMNH 33305, Lectotype, Indonesia. 52.8 mm SL, by T Fraser. D. BPBM 27640, 45 mm SL, Kovalam, India, by J. Randall. E. USNM 276669, 49.2 SL, Pulau Nias, Sumatra, Indonesia, by T. Fraser. Taeniamia leai F–K: F. Adult, near Noumea, New Caledonia, by R. Myers. G. Adult, Lord Howe I., by N. Coleman. H. Juvenile, New Caledonia, by G.R. Allen. I. USNM 356269, 55 mm SL, One Tree I., Queensland, Australia. J. F308, Syntype, 68 mm SL, Norfolk I., by T. Fraser. K. Figure of Archamia leai, syntype, 68 mm SL, by E. R. Waite, 1916 from Plate 45.

FIGURE 2. Radiographs as positives. A. Apogon bleekeri, RMNH 33905, lectotype, 52.8 mm SL, Indonesia by D. Smith. B. Archamia leai, USNM 356272, 53 mm SL, One Tree I., Australia by S Raredon. C. Holapogon maximus, BMNH 1889.4.15.22, 179 mm SL, Muscat, Oman, by T. Fraser. Insets of each caudal skeleton. Hypurals = 1–5, terminal centrum = tc, tenth vertebrae = Ą, length of anterior supernumerary spine = [.

FIGURE 3. Anterior and mid-dorsal fin elements. A & D. Archamia bleekeri, USNM 356291, 37.6 mm SL. B &E. Taeniamia leai, USNM 306823, 55.0 mm SL, bone stain only. C & F. Kurtus indicus USNM 267150, 63.0 mm SL. supraneurals = S1–3; neural spines = 1–3, 6–9; proximal-middle radial = P1–2, 5–8; distal radial = R; fused radials = FR; dorsal spines = I–III V– VII and IX; shaded areas = cartilage stain. Scale = 1 mm.

FIGURE 4. Radiographs as positives of the lower mid-section of the body and anterior anal fin elements. A. Archamia bleekeri, NTM S.13958.020 by H. Larson. B. Taeniamia leai, USNM 356277 by Sandra Raredon. C. Kurtus indicus, USNM 372503 by Sandra Raredon. D. Holapogon maximus, BMNH 1889.4.15.22 by T. Fraser. tenth vertebrae = 10, flange = f, ribs = r4–12, proximal-middle radials = mp1–5, nubbin = n, supernumerary spines = S, = serial spine with respect to anteriormost proximal-middle radial = I, homologous positions with respect to serial proximal-middle radials 1 or 2 = s1–2.

FIGURE 5. Lateral views of vertebrae 9–12 with haemal arches and associated ribs, haemal spines and proximal portions of the anal fin pterygiophores. A. Taeniamia macroptera, USNM 171228, 54.0 mm SL. B. Archamia bleekeri, USNM 356291, 37.6 mm SL. C. Kurtus indicus, USNM 267150, 63.0 mm SL. = tenth vertebra = 10, = haemal arch = HA, = haemal spine = HS, ribs = R8−10, inferior foramen = IF, tips of anal fin proximal radials = 1–10, shaded areas = cartilage stain. Scale = 1 mm.

FIGURE 6. Last few fin elements of the dorsal and anal fins. A & E. Taeniamia leai, USNM 306823, 55.0 mm SL, bone stain only. B & F. Archamia bleekeri, USNM 356291, 37.6 mm SL. C & G. Kurtus indicus USNM 267150, 63.0 mm SL. D & H. Kurtus gulliveri, USNM 217309, ~50 mm SL. d = distal radial; fr = fin ray, m = middle radial, p = proximal radial, p+m = cartilage not free from proximal radial, shaded areas = cartilage stain. Scale = 1 mm.

FIGURE 7. A. Archamia bleekeri, USNM 356291, 37.6 mm SL, anterior eleven vertebra, ribs and epineurals. B. Taeniamia leai, USNM 306823, 55.0 mm SL, anterior eleven vertebra, ribs and epineurals (not counter stained). C. Taeniamia leai, USMN 306823, 55 mm SL, partial portions of ribs 1–4 enlarged to show narrow flanges on ribs 2–4. D. Kurtus indicus, USNM 267150, 63.0 mm SL, vertebra 3–6, ribs and epineurals (starting on v4 and ending on v6). Epineurals in red and cartilage in blue. Vertebra number on centrum = 1, 3, 5, 11, rib number = R3. Scale = 1 mm.

FIGURE 8. Caudal fin elements from third pre-ural centrum. A. Archamia bleekeri, USNM 356291, 37.6 mm SL. B. Taeniamia leai, USNM 306823, 55.0 mm SL, only cleared and stained. C. Kurtus indicus, USNM 267150, 63.0 mm SL. D. Kurtus gulliveri, USNM 217309, ~50 mm SL. Numbers refer to upper and lower procurrent (5–14) and principle caudal-fin rays (8–9). terminal centrum = ct, epural = e, = hypural = h, neural crest = nc, = neural spine = ns, parhypural = p, preural centrum = pu, urostylar sheath = us = paired uroneurals = u, fused elements = +, paired uroneurals in solid black, cartilage in gray. Scale = 1 mm.

FIGURE 10. Dorsal view of the upper branchial bones, all similarly sized. A. Apogon townsendi AMNH 23513, modified from figure 41B, Rosen and Patterson (1990). B. Taeniamia leai USNM 306823, 55 mm SL, bone stain only. C. Archamia bleekeri, USNM 356291, 37.6 mm SL. D. Kurtus gulliveri, AMNH 43396, modified from figure 46A, Rosen and Patterson (1990). E. Kurtus indicus USNM 267150, 63.0 mm SL. F. Taeniamia buruensis USNM 112112, 62.0 mm SL. second epibranchial = E2, interarcual cartilage = IC, third pharyngobranchial = P3, shaded areas = cartilage stain.

FIGURE 11. Hyoid series. A. Archamia bleekeri, USNM 356291, 37.6 mm SL, outer right side laterally reversed. B. Kurtus indicus, USNM 267150, 63 mm SL, outer left side. upper hypohyal = UH, lower hypohyal = LH, branchiostegals = BR1–7, = beryciform foramen = BF, anterior ceratohyal = AC, posterior ceratohyal = PC, interhyal = IH. Scale = 1 mm.

FIGURE 12. Dorsal view of infraorbitals with shelves. A. Archamia bleekeri, USNM 356291, 37.6 mm SL, right side. B. Taeniamia leai, USNM306823, 55 mm SL, left side laterally reversed. C. Kurtus indicus, USNM 267150, 63 mm SL, right side. infraorbital = 1–6, shelf = S. Scale = 1 mm.

FIGURE 14. Dorsal (A) and lateral (B) view of Taeniamia leai, USNM 306823, 55 mm SL.

FIGURE 15. Dorsal (A) and lateral (B) view of Taeniamia macroptera, 53 mm SL from Fraser (1972) plate 37. Scale = 1mm.

FIGURE 16. Semi-diagrammatic fourth lateral-line scales, left side, growth rings not shown. A. Taeniamia leai, USNM 356272, 53 mm SL. B. Archamia bleekeri, USNM 276669, 50 mm SL. C. Kurtus indicus UF 182306, 94 mm SL, scale enlarged 7.6X. D. Kurtus indicus UF 182306, 94 mm SL, lateral-line scale at size for size comparison with scale illustrated for Archamia bleekeri. Scale for A, B and D = 1 mm., Scale for C = 0.25 mm.

FIGURE 17. Partial dorsal view of pores and free neuromasts (dots). Scales = 1 mm. A. Taeniamia leai, semi-diagrammatic, free neuromasts (dots) obscured or missing, USNM 356269, 65.1 mm SL. B. Archamia bleekeri, semi-diagrammatic, some pores and free neuromasts (dots) obscured or missing on head, USNM 276669, 50.0 mm SL.

FIGURE 18. Partial lateral views of the head. Scale = 1 mm. A. Taeniamia leai, semi-diagrammatic for pores, free neuromasts obscured or missing, pores missing on cheek, USNM 356269, 65.1 mm SL. B. Archamia bleekeri, semi-diagrammatic, pores and free neuromasts (dots) obscured or missing on part of cheek and behind eye, USNM 276669, 50.0 mm SL.

FIGURE 19. Ventral view of left dentary from mental to articular pores. Scale = 1 mm. A. Taeniamia leai, semi-diagrammatic for pores, free neuromasts obscured or missing, USNM 356269, 65.1 mm SL. B. Archamia bleekeri, semi-diagrammatic, for pores, free neuromasts (dots), dentition of premaxilla partially visible. USNM 276669, 50.0 mm SL.