Foa

Fraser, Thomas H. & Randall, John E., 2011, Two new species of Foa (Apogonidae) from the Pacific Plate, with redescriptions of Foa brachygramma and Foa fo, Zootaxa 2988, pp. 1-27 : 10-15

Foa fo Jordan and Seale, 1905

Figure 6–8 , 14–15 , table 1–4.

Material examined: Lectotype Foa fo CAS SU 9672 metal tag, 41.0 mm SL, ~52.3 mm TL, Philippines, Luzon I., Cavite, 1901, G. A. Lung.

Other material. Australia: AMNH 33738 (3, 15-19), Queensland, Endeavour reef, 7 Sep 1969. AMS I.27783.003 (6, 18-37), Queensland, First Lagoon, 23.1°S 152.1°E, 3–4 m, 24 Nov 1987. Indonesia: BPBM 37336 (1,), Flores I., Maumere, off mouth of creek, 17 Nov 1996. USNM 261582 (1, 24), Sulawesi Ids., Kabaena I., Tallabassi Bay, 5°17'20"S 122°04'E, VGS-74- 2, 2–15 m, 25 Feb 1974. Taiwan: SU 20996 (1, 46), Kaohsiung (Takao). Philippines: CAS SU 9672 (1, 45.5) 55.2 mm TL, Luzon I., Cavite, 1901, G. A. Lung. SU 20266 (1, 36), Iloilo, Panay I. USNM 56279 (1, 37.5), Cavite, x-ray. USNM 169162 (31, 15-32), Iloilo, (1, 33) cleared and stained. USNM 171102 (2, 33-36), Albatross, 20 May 1908, cleared and stained. USNM 268164 (11, 17-25), Negros I., Canauay river, 09°3'N 123°03'48"E, LK 79-12, 0–0.9 m, 9 May 1979. USNM 344871, (39, 16.6-33.2), Negros, Dumaguete Market, 27 Mar 1976. Palau, CAS 83099 (2, 27-27), Urukthapel I., 07°16'18 "N 134°26'43 "E, GVF 0 483, 2.5 m, 12 Sep1954. Saipan: USNM 154270 (1, 34.1), West Lagoon, 3 Apr 1949. Guam: UGM 6658 (1, 29), Tumon Bay, photograph. Caroline Islands: USNM 235047 (3, 19-34) Ponape, South of Nanmatol I., 06°50'45"N 158°19'00"E, RC-3, 0–. 5 m, 4 Jul 1981. Papua New Guinea: USNM 262582 (5, 23-37), Trobriand Is., Kiriwina I., 08°31'48" S 150°59' 36"E, BBC 1694, 0–0.6 m, 16 Jun 1979. USNM 370287 (12, 18.8-36.0), Taurama Pt, SE of Port Moresby, BBC 1527, 20 Jun1970. New Caledonia: BPBM 28668 (1, 28), Noumea, channel beween Ile Canards and Ile Maitze, 20 m, 19 Mar 1975. SMNS 19730 (1, 28), Grande Terre, Bale de Port-Bouquet, 21°42'35"S 166°23'57"E, 0.5– 2 m, 8 Nov 1997. SMNS 21557 (6, 21-38), Grande Terre, Enghoue, 22°09'14"S 166°14'33"E, 0–0.1 m, 13 July 1999. 13 July 1999. SMNS 22854 (2, 21-30), Ile des Pins, 22°34'05"S 167°25'23"E, 0–2.5 m, 16 May 2000. SMNS 22997 (3, 19-27), Grande Terre, Baie de Pritzbue, at Bourake, 21°57'32"S 165°59'20"E, 0–2.5 m, 23 May 2000. USNM 268244 (2, 22-24), Grande Terre, Chenal De L'Ilot Maitre, 18– 25 m. Solomon Islands: USNM 358155 (1, 23), Santa Cruz Islands, Duff Islands, Taumako Island, 09.9° 167.1583°W, SOL 98-21, 0.5m, 22 Sep 1998. Fiji: USNM 262588 (16, 26-37), Matuku I, main harbor, 19°09'38"S 179°45'23"E, VGS 82-3, 23 Apr 1982. USNM 262592 (1, 22), Naviti I., 17°06'S 177°13'E, VGS 82-32, 28 May 1982. Samoa: AMNH 58010 (4, 15-32), Tutuila I., Pala lagoon near center, PH-72-II-29, 29 Feb 1972. USMN 51735 (1, 26.7), Upolu, Apia, 1902, D. Jordan and V. Kellogg, x-ray (in poor condition). USNM 220060 (2, 24-25), Tutuila I., Fatu Mufuti, 0–36 m, 1977. USNM 325625 (2, 21-33), Western Samoa on Reef Flat near Apia, 0–2 m, 24 Mar 1979. Wallis Islands: USNM 376768 (1, 27), Ile Uvea, on reef flat off NW end of island, just off Afe Utu'uhu, 13.2181°S 176.219°E, JTW 2000- 17, 1m, 11 Nov 2000. USNM 376748 (4, 31-37 mutilated), Ile Uvea, outside barrier reef at NW end of island, 13.2097°S 176.246°E, JTW 2000- 16, 22m, 11 Nov 2000. Tonga Islands: BPBM 38308, (2, 44.1-53.5), Tonga, Mar 1983. French Polynesia: CAS 83106 (23, 17-42), Tahiti, Taravao, lagoon, GVF 1104, 1 m, 21Apr1956.

Diagnosis. Pored lateral-line scales usually 8–12; gill rakers and rudiments usually 13–15; body scales margins uniform to pale edging; body mottled brownish with five darker irregular bars, first anterior to origin of first dorsal fin, second from posterior half of first dorsal fin through posterior half of pelvic fin, third narrow bar between soft dorsal and soft anal fins, fourth on caudal peduncle, fifth at base of caudal fin; whitish spot on axil of pectoral fin; three whitish basicaudal spots partially in darkish basicaudal bar; whitish spot behind posterior base of soft dorsal fin; second dorsal fin, anal fin and caudal fin pale with darkish banding.

Description. Range of proportions as percent of standard length in Table 1 . The lectotype is described and variation is given in Tables 2–4.

Dorsal fin IX spines as VII(I)-I,9, third spine longest and strongest, sixth radial free in advance of hidden nubbin representing eighth spine; anal fin II,8; pectoral-fin-rays 12; pelvic fin I,5; principal caudal rays 1-8+7+1; lateral-line scales 23 with 14 pored and 9 partial pores and pits ( Table 4); transverse scale rows above lateral line 1; transverse scale rows below lateral line 4; median predorsal scales 4; circumpeduncular scale rows 12 as 5+2+5; gill rakers + rudiments 14 ( Table 3), well- developed rakers 8, upper arch with 2 rudiments and 1 raker, lower arch with 7 rakers and 4 rudiments; second arch with rudiments on hypobranchial and ceratobranchial with one raker at angle.

Villiform teeth in a band on the premaxilla and dentary; 2–3 rows on vomer; 3 rows on the palatine; none on ectopterygoid, endopterygoid or basihyal.

Vertebrae 10+14; five free hypurals; one pair of short, slender uroneurals; three epurals, the first two expanded, a free parhypural; three supraneurals, no procumbent spines (spurs); two supernumerary spines on first dorsal pterygiophore, no procumbent spines (spurs); basisphenoid reduced to upper portion (meningost); supramaxilla thin and reduced in length; posttemporal smooth on posterior margin; preopercle smooth on vertical and horizontal edges, ridge smooth; infraorbitals smooth, infraorbital shelf present on third bone; interhaemal gap 1+4 (Table 2).

Peripheral ctenoid scales on cheek, subopercle, opercle, isthmus, base of pectoral fin, ctenoid behind pectoral fin and on body, nape and predorsal scales ctenoid; two large cycloid scales on base of pelvic fin, no axillary scale; cycloid scales onto caudal fin about same size as scales on caudal peduncle.

Head of lectotype with skin congealed, obscuring pore openings and without evidence of sensory neuromasts; anterior end of supra-orbital canal as a broad slit at edge of upper lip curving up distally; lachrymal with large anterior pore near flat posterior nare opening, two large ventral slit pores along edge of lachrymal; anterior portion of dentary with dentary (anterior) and mental (ventral) pores, posterior with a large articular pore; supratemporal canal with posterior canals onto first nape and predorsal scales.

Head of lectotype with most free neuromasts missing. Free neuromasts on snout, interorbit and temporal areas in ( Fig. 7 A), a split line becoming one line of neuromasts from snout interrupted over eye, offset, not overlapping, continuing onto side of head below posttemporal, short lines generally perpendicular to long lines, short medial lines associated with commissure region, supratemporal neuromasts missing; several rows of neuromasts between nares and anterior infraorbital oriented horizontally and vertically; a long linear line of neuromasts from first infraorbital (lachrymal) past posterior corner of maxilla dividing on to lower cheek horizontally as well as downward, short lines of neuromasts radiate from ventral and posterior edge of eye ( Fig. 7 B); three or four rows of neuromasts on anterior third of lower jaw grading posteriorly to one row near edge of articular pore ( Fig. 7 C), gular free neuromasts in chevron rows except for anterior two rows; preopercle with single long vertical line of neuromasts on upper arm from posterior part of lower arm (7C & 8A); preopercle flap (horizontal arm) with short rows of various orientations; ( Fig. 8 A); opercle with short vertical row of neuromasts on upper anterior scale ( Fig. 8 A), a long vertical row near preopercular edge, a row on posterior scale, and two areas with neuromasts on opercular flap; pored lateral-line scales with vertical rows of neuromasts above and below canal pore;, first scale with pit with a reduced row above pit, free neuromasts on other pit scales and several small caudal scales, no free neuromasts observed on other basicaudal scales ( Fig. 8 B); free neuromasts associated with 11 principal caudal fin-rays in single linear lines on ventral edges of upper branched rays (3–8) and on dorsal edges of lower branched rays (11–15), middle two caudal rays (9–10), upper unbranched and branched caudal rays (1–2), and lower unbranched and branched caudal rays (16–17) lack free neuromasts as in fig. 5B.

Anterior nare tubular, posterior nare flat.

Caudal fin rounded, broken in lectotype; second dorsal and anal fin with rounded distal edges.

Life colors. Figure 6 A. Head, body, first dorsal and pelvic fins brown with white spots and markings; darker brown bar-like markings on head and body, a cheek mark from eye to upper jaw, a mark above preopercle over anterior nape, one or two short marks above opercle not reaching base of first dorsal fin, an irregular bar from near mid-base of first dorsal fin to side of abdomen, an irregular bar from base of soft anal rays to base of soft dorsal rays, an anterior caudal peduncle mark, and diffuse basicaudal marks; lips brownish with narrow alternating white marks; edge of eye with alternating brown and white markings; irregular whitish spots on body and caudal peduncle, a white spot at upper pectoral fin base, a white spot at mid-base of soft dorsal rays, a white spot and posterior base of soft dorsal fin, three basicaudal white spots; membranes between second to fourth dorsal spine more whitish than brownish, fourth to seventh membranes more brownish than whitish; base of pelvic rays to tip of spine whitish, remainder of fin brownish; soft dorsal, anal and caudal fin rays with tiny white spots on translucent fins. See Kuiter and Kozawa (2001, p. 95, figs. A–C & E) for color variations from Indonesia.

Post mortem color pattern. Figure 6 B. Head and body tan with brown markings, whitish spots in life mostly not apparent except for pale area at posterior base of second dorsal fin and three pale areas on base of caudal fin outlined by two darker spots; diffuse cheek mark; faint markings on body consistent with dark markings in life; first dorsal fin dusky with mid-paler region; second dorsal fin with brownish base and tiny dark markings on rays and pale membranes; pelvic, anal and caudal fin rays with tiny dark markings on rays and pale membranes

Preserved color pattern. The lectotype has no markings on head, body or fins. Other material is consistent the description by Jordan and Seale (1905) of Foa fo from the Philippines as follows: “Color in spirits, yellowishwhite, mottled with brownish, the mottlings assuming a more or less irregular band-like arrangement over back and sides; a dark stripe from lower posterior margin of orbit to lower limb of preopercle; a dark band over posterior part of nuchal region; about 5 very indistinct darker bands around under part of head; spinous dorsal with two dusky blotches; soft dorsal yellowish; anal yellowish with about 5 indistinct brownish lines; ventrals with inner half and posterior part deep black; pectorals yellow; caudal yellow with four indistinct cross-bands of dusky; iris golden, darker above”.

Figure 6 D is from Jordan and Seale (1906) who described Foa fo from Samoa as follows:

“Color in spirits, yellowish white, the scales shaded more or less with light brown; five diffuse, irregular vertical cross-bands of dusky, the first in front of dorsal, the second from middle of spinous dorsal, the third from base of soft dorsal, the two posterior ones encircling caudal peduncle; there is also a more or less distinct dusky area on nuchal region; dusky cross-shades under chin; the fins, except pectorals, all more or less indistinctly blotched with dusky, the spinous dorsal being black at tip with a lighter area in middle and dusky at base; anal shows indications of three or four darker cross-shades; ventral dusky; a dark spot on opercle, just posterior to eye; upper portion of orbital and snout shaded with dusky”. Figure 6 C show a typical color pattern of a tannish background with darker markings generally consistent in location with post mortem and live color patterns; an additional post ocular mark is more apparent extending from eye to near origin of the lateral line scales.

Distribution. Widespread, found on most Pacific Plate islands out to French Polynesia, islands bordering the Pacific Plate, Australia, New Guinea, Philippines ( Fig 14 ). Although we did not report on material from the Indian Ocean, Gon and Randall (2003) reported from East Africa and the Red Sea.

Remarks. We regard color descriptions in 1905 and 1906 by Jordan and Seale as being fairly consistent and without significant variances that might suggest two different species. Color photographs suggest that another species of Foa may be present in the Philippines,http://fishpix.kahaku.go.jp/fishimage-e/index.html #KPM- NR0035772 from Cebu I. The location of the syntype of Foa fo from Negros, Philippines is unknown. There were two specimens in CAS SU 9672 from Cavite. Lengths were not given in the original description for the Cavite specimen. We chose one. There are several live photographs (#945135-37, 39; 954289; 95911-12) of Foa fo from Leyte and Bohol, Philippines (http://www.poppe-images.com/index.php?t=1).

Gon and Randall (2003) also discussed the connected history of Foa fo and F. brachygramma as well as the false type from Samoa identifying F. f o from the Red Sea. The fish from Inhaca, Mozambique in their plate 5I may represent a different species, most likely Foa madagascarensis Petit, 1931 , given his color description: “ Coloration (en alcool). -— Couleur générale brun chamois, plus claire sur le ventre. Tout le corps marqu d'un pointill de petites taches brunes, plus grosses et plus espacées sur les écailles du ventre. Membrane unissant les rayons de la première dorsale de couleur foncée (fum fonc). Celle qui unit les rayons de la seconde dorsale, plus claire. Les rayons branchus de la ventrale ont une extrémit claire, suivie d'une longue zone noire. La base est claire. Pectorale et caudale claires”. “Ground color brown yellow, paler on abdomen. All of the body marked with linear small brown spots becoming larger and farther apart on abdominal scales. First dorsal fin membranes darkish. Membranes of the second dorsal fin lighter than in the first dorsal fin. The branched rays of the pelvic fin have pale tips and distally black except for the base which is pale. Pectoral and caudal fins pale”. Specimens of Foa from Madagascar (USMN 203771), South Africa (ANSP 53447) and Tanzania (USNM 344874) are consistent with Petit’s color description about having small brown spots. No such color description was given for the Red Sea specimens provisionally identified by Gon and Randall (2003) but their description of bars on the body and caudal peduncle are suggestive of Foa fo. An underwater photograph by M. Hackenberg from the Red Sea ( Lieske and Myers, 2004:90 misidentified as Fowleria vaiulae ) has the whitish spots and irregular brown bars on a light brown ground color as seen on Foa fo as do underwater photographs from J. L. Rose at Sharm el Sheik in the northern Red Sea ( Fig. 6 c). Allen in Randall and Lim (2000) listed Foa madagascarensis from the South China Sea, likely one of the other species.

Figures

FIGURE 6. Foa fo: A. Live from Palau by Hirashi Nagano. B. Post mortem USNM 28.7 mm SL, Wallis I, by J. T. Williams. C. Live from Sharm el Shiekh, Egypt by Jean Louis Rose. D. False type of Foa fo, USNM 51735, 27 mm SL, Samoa, Upolu, Apia, in Jordan and Seale (1906), digital scan of original published as fig. 42, drawn by Chloe Starks.

FIGURE 7. Foa fo, AMS I. 27783 - 003, 36 mm SL. semi-diagrammatic views of the head. A. dorsal. B. right side reversed. C. lower jaw, right side, reversed. pores = shaded; free neuromasts (dots).

FIGURE 8. Foa fo. A. AMS I. 27783 - 003, semi-diagrammatic views opercular region with pores (shaded) and free neuromasts (dots). B. AMS I. 27783 - 003, 28.9 mm SL, scales at the base of the caudal fin with ctenoid and cycloid scales, last pitted lateralline scale, and free neuromast (dots) patterns on scales. Right side laterally reversed.

FIGURE 14. Distribution of collection sites. Foa brachygramma =, F. f o =, F. n i v os a =, F. leisi =.

FIGURE 15. Variation in the number of pored lateral-line scales with size in four species of Foa. F. brachygramma =, F. f o =, F. n i v o s a =, F. leisi =.