Apogonichthyoides Smith, 1949

Fraser, Thomas H. & Allen, Gerald R., 2010, Cardinalfish of the genus Apogonichthyoides Smith, 1949 (Apogonidae) with a description of a new species from the West-Pacific region, Zootaxa 2348, pp. 40-56 : 41-45

Apogonichthyoides Smith, 1949

Figure 1 A–C

type species: Amia uninotata Smith and Radcliffe in Radcliffe, 1912

Diagnosis: Species with IX dorsal spines as VIII-I or VII(I)-I with eighth spine reduced to a tiny visible spine or a nubbin hidden under skin supported by a free sixth distal radial, third spine the longest and thickest; 9–10 soft dorsal rays, fin edges rounded; II anal spines, second longest; 8–9 soft anal rays, fin edges rounded; 14– 17 pectoral-fin rays; longest procurrent caudal rays segmented, 1+8-7+1 principal caudal rays; emarginate to truncate-caudal fin; 14–17 total rudiments and rakers on first gill arch; 7–12 well-developed gill rakers, small tooth patches or low rudiments present on second ceratobranchial and hypobranchial; three supraneurals, no procumbent spines (spurs); two supernumerary spines on first dorsal pterygiophore, no procumbent spine (spur) on first dorsal pterygiophore; one supernumerary spine on first anal pterygiophore; interhemal gap 1+3 or 1+4; one pair of slender uroneurals with slight anterior expansion; three epurals, largest with anterior expansion; free parhypural, five free hypurals, two autogenous haemal spines; cheek, subopercle, opercle, nape and body scales ctenoid; 1–3 predorsal scales cycloid or ctenoid, two large median pelvic scales cycloid or ctenoid, no visible developed accessory pelvic scale (may be hidden by large scale in some species); pored lateral-line scales with two or three openings above and one or two below lateralis canal; last lateral-line scale slightly elongated to elongate with one to three small openings laterally; cephalic lateralis pores numerous, mostly as minipores; lachrymal with a large terminal pore and a pair of large ventral pores; dentary and mental pores present; terminal end of supraorbital canal with a large slit pore at edge of snout; margins of supraorbital canal with small pores to intersection with supratemporal canal; infraorbital canal with a few projections and minipores over cheek scales; smooth anterior margin of supratemporal canal, well-developed posterior supratemporal canal margin with complex projections and minipores; lateral margins of mandibular canal with many minipores; a small posterior articular pore; anterior nare tubular, posterior nare opening flat; stomach and intestine usually pale or rarely blackish; pale peritoneum; no dark or light stripes on snout, through eye or continuing posterior to eye; no midline body stripe contiguous or in line with eye; head usually with darkish cheek mark; darkish markings posterior of eye, on nape or associated with the posttemporal; two faint to intense darkish bars on body associated with each dorsal fin; ocellus or ocellus-like mark in anterior bar of some species; bars may be obvious only in young or juveniles of some species; faint to intense darkish basicaudal bar, some with intense spot in basicaudal bar or with dark basicaudal spot; some species with faint or without body or peduncular bars but with irregular to regular darkish spots on body; no stripe in anal fin; stripe-like darkish marks in second-dorsal fin or without a stripe; pelvic fins with a whitish outer edge along spine or with a mottled whitish to yellowish area extending from base of the fin.

Remarks: Our recognition of Apogonichthyoides , formerly placed in the synonymy of Apogon is based on color patterns observed in juveniles and adults, morphological characteristics and emerging molecular hypotheses. In the following paragraphs we provide historical background information for this reconstituted grouping, of which the wide-ranging Apogon timorensis is a better-known species. See additional remarks under each species description.

Smith created Apogonichthyoides in 1949. His type species, Amia uninotata Smith and Radcliffe in Radcliffe, 1912, was from the Philippines. Smith illustrated what he identified as the type species but was likely a specimen of Apogonichthyoides pharaonis ( Bellotti, 1874) , a species known from the Red Sea, East Africa and the Arabian Sea ( Gon, 2000). Smith (1961) believed this genus had ‘... an admittedly tenuous taxonomic basis but it is a convenient method of grouping the species... ’. We provide figures 1A–C to portray Smith’s type species from the original illustration of the type, a postmortem specimen and an underwater photograph. This species is known from seven collections in the Philippines.

Smith (1961) described Apogonichthyoides enigmaticus from a specimen in poor condition (fins), noting the good condition of the body, darker edging of body scale margins and villiform bands of teeth in each jaw, some on the side of dentary slightly enlarged. Gon (1993) concluded that this specimen was Apogon apogonides ( Bleeker, 1856) after a detailed analysis. The interpretations of Smith’s observations given above were discounted rationally. However, Apogon dhofar described by Mees in Randall (1995, figure 394) and by Mees (1996, figures 1, 3–4) clearly show the pattern of darker edging on the body scales. Data taken from Smith’s type, Mees’ type material and specimens from Somalia (USNM 213325 and USNM 304739) by the first author were in good agreement for dentition, gill raker and pectoral-fin ray counts. We believe Smith’s species is valid and Apogon dhofar is a junior synonym. The case for including this species in Apogonichthyoides is not clear. Juveniles have two faint, narrow bars on the body and a small caudal peduncle spot in juveniles both fading in adults. White-edged pelvic fin-spines of juveniles and adults along with the juvenile color pattern suggest inclusion. Slightly higher well-developed gill raker counts, a tendency for elongated anterior soft dorsal rays along with the color pattern suggested potential relationships with Apogon griffini and Apogon flagelliferus . We provisionally leave Smith’s species in his genus.

Fraser (1972) placed Apogonichthyoides niger (Döderlein in Steindachner and Döderlein, 1883), A. opercularis , A. taeniatus (Cuvier in Cuvier and Valenciennes, 1828) and A. timorensis , in the species rich catch-all subgenus Nectamia , later subsumed in the subgenus Ostorhinchus by Gon (1987). Randall (2005) recognized Ostorhinchus as a genus largely based on Bergman’s (2004) unpublished PhD dissertation treating apogonid relationships. Her representatives of Ostorhinchus were few, examining sensory pore patterns of just five species (including A. timorensis ).

Gon and Randall (2003) compared Apogon timorensis with A. pharaonis and other species in the nigripinnis complex ( Gon, 2000). The juvenile color patterns of Apogon niger and A. pharaonis illustrated by Gon (2000) are similar to the color pattern for the new species described below. Both the juvenile and adult color patterns of Apogon niger (see Kuiter and Kozawa, 2001) have common pattern features with the timorensis lineage. The general color pattern similarity with darkish bars and spots, a well-developed nape lateralis pore system, and low number of developed gill rakers suggests that Apogonichthyoides includes members of both the nigripinnis and timorensis complexes.

Mabuchi et al., (2005) provided molecular evidence using 1287 base pairs from mitochondrial genes 12S rRNA, 16S rRNA and tRNA val that Apogon sensu lato was not monophyletic. They recognized three major groups within Ostorhinchus . Their work showed that color patterns were associated with phyletic lineages and had ancient origins within the family. Their results suggested more species needed to be included in analyses now being pursued by Mabuchi, Fraser, Azuma and Nishida. One major group (Group II of Mabuchi et al., 2005) includes the members of the genus Nectamia recently recognized by Fraser (2008). Representatives of Apogonichthyoides were present in Group II (barred pattern) but not with representatives of Ostorhinchus in group III (striped pattern). Furthermore, Apogonichthyoides differs from Ostorhinchus by lacking a snout/ head stripe, possessing bars or darkish spots on the body instead of having a uniform body or one or more stripes, a weakly emarginate to truncated caudal fin instead of forked, larger/longer pelvic fins, tubular anterior nare instead of a low rim with a flap, a small rudimentary eighth dorsal spine instead of absent and a free sixth radial instead of fused.

Mabuchi et al. (2005) reported Apogon ishigakiensis Ida & Moyer, 1974 as part of their Group II. This species is similar to or possibly synonymous with Apogon hoevenii Bleeker, 1854 c. The provisional tree recovered in July 2009 places both of these species in a clade elsewhere with Apogon rueppllii Günther, 1859 (Mabuchi, Fraser, Azuma and Nishida, unpublished data, American Society of Ichthyologists and Herpetologists annual meeting, 2009). The color pattern of larval and juvenile Apogon rueppllii have no bars on the body ( Neira, 1991). This species is not treated as a member of Apogonichthyoides . We do not include Apogon hoevenii or A. ishigakiensis in the timorensis complex of Apogonichthyoides because their lower pectoral fin-ray counts (12) and higher well developed gill-raker counts (14–16). Mabuchi et al. (2005) included another brownish-black species, Apogon melas Bleeker, 1848 in Group II, but without support for a close relationship with Apogonichthyoides .

Fraser (2006) identified Vincentia and several species of uncertain relationship that may be related to the timorensis group, to which we add Quinca mirifica Mees, 1966 based on adult and juvenile color patterns. Bergman’s (2004) observation the cephalic lateralis pore pattern of Apogon timorensis was similar in most respects to Vincentia chrysura ( Ogilby, 1889) , however, this species is not the type of Vincentia as she stated. We place chrysura in Apogonichthyoides based on the cephalic lateralis pore pattern and general color pattern. Vincentia was recovered in a clade with Apogonichthyoides as part of a provisional molecular tree well separated from Ostorhinchus (Mabuchi, Fraser, Azuma and Nishida, unpublished data, American Society of Ichthyologists and Herpetologists annual meeting, 2009).

Apogonichthyoides include the A. timorensis complex with eight species and A. nigripinnis complex with 11 species. All names are new combinations except Apogonichthyoides enigmaticus , A. nigripinnis , A. taeniatus and A. uninotatus used by Smith (1961).

Included species: nigripinnis group Apogonichthyoides atripes ( Ogilby, 1916) , A. cantoris , A. enigmaticus , A. heptastygma (Cuvier in Cuvier and Valenciennes, 1828), A. niger , A. nigripinnis (Cuvier in Cuvier and Valenciennes, 1828), A. pharaonis , A. pseudotaeniatus, ( Gon, 1986) , A. sialis ( Jordan & Thompson, 1914), A. taeniatus , and A. uninotatus . timorensis group Apogonichthyoides brevicaudatus ( Weber, 1909) , A. chrysurus , A. euspilotus ( Fraser, 2006) , A. gardineri , A. opercularis , A. regani ( Whitley, 1951) , A. timorensis , A. umbratilis new species.

Material examined other than listed in species accounts: Apogon atripes . NMV 3175, (1, 59.9), Australia, Queensland, 12°03'17"S 143°15'10"E. NMV R5961, (1, 71.9), Australia, Queensland, 19°58'S 148°41'E. Apogon brevicaudatus Syntypes ZMA 101.127, (2, 35.4–100.7) Indonesia, Aru Is., Jedan I. Apogon bifasciatus Syntypes SMF 2186, 4622, 4623 (3, 63.1–67.7), Red Sea, Rüppell. Syntypes Apogon cantoris RMNH 5599 (2, 83.2–84.9), Rio(Rhiau Islands, near Singapore) Indonesia. Apogon chrysurus Syntypes I. 1799-1801, (3, 51.0–54.9), Australia, Lord Howe Island, 1888. USNM 213338, 3, 31–38, Australia, One Tree I., 11 Dec 1966, x-ray. Apogon dhofar Holotype CAS 82327 (1, 78.9), Oman, 20 Feb 1990, x-ray. Paratypes BMNH 1995.9.20, 2, 52.5–55.6, Oman, 20 Feb 1990, 45– 80 m. Apogonichthyoides enigmaticus Holotype SAM 13851 (1, 79.6), South Africa, Durban, 1915, x-ray. USNM 213325 (6, 58–76), Somalia, 11°14'N 51°08'E, Anton Bruun, Cr9, Sta 456, 17 Dec 1964. USNM 304739 (1, distorted), Somalia, 10°39'12"N 51°18'48"E, RV Bienta, 16 Oct 1985, 46 m. Apogon enneastigma Syntypes SMF 1065, 12130 (3, 32.6–37.0), Red Sea, Rüppell. Apogon euspilotus Holotype CAS 118836 (1, 59.5), New Caledonia, Nouméa, Ile aux Canards. Apogon heptastygma USNM 213355 (32, 22.8–47.6), Eritrea, Sheikh El Abu, 16°08'N 39°26'30"E, 14 Aug 1969. Apogon hoevenii Holotype RMNH 5581 (1, 40.2), Indonesia, Amboine. SU 27370 (4, 43.5–50.2), Philippine Is., Dumaguete, 27 Jun 1931. SU 33473 (60, 17.7–51.4), Philippine Is., Dumaguete, 21 Dec 1936. Apogon ishigakiensis Holotype ZUMT 52989 (1, 37.4), Ryukyu Is., Iriomote-jima. Apogon niger CAS 52217 (1, 50.0), Philippines, Siluag I. 23 Jun 1948. BMNH 1891.5.26.4 (1, 68.2), Japan, Tokyo Bay. Apogon nigripinnis Syntype MNHN 8694 (1, 63.4), India, Pondicherry, x-ray. Apogon pharaonis SAIAB 3149 (2, 60–76), Mozambique, x-ray. SAIAB 3151 (1, 67), Mozambique, 1938, x-ray. SAIAB 3148 (4, 42– 84), Mozambique, x-ray. SAIAB 3150 (4, 68–74), Mozambique, x-ray. SAIAB 3152, (1, 69), Palma, 1 Aug 1951, x-ray. SAIAB 3153 (2, 49–53), Aldabra I., 14 Nov 1954, x-ray. Apogon pseudotaeniatus Holotype BPBM 27395 (1, 87.2), Egypt, Gulf of Aquba, 16 May 1969. Apogon punctatus Syntypes BMNH 1908.3.23.86–89 (4, 31.9–65.4), Cargados Carajos (St. Brandon Shoals). Apogon rueppellii Syntype BMNH 1972.6.8.2, (1, 55.5), Australia. Amia sialis Holotype FMNH 57084 (CM 6021) (1, 90.4), Japan, Surugu Gulf. Apogon suezii Holotype MNHN 5137 (1, 50.6), Egypt, Suez, Letourneux. Apogon taeniatus Syntypes MNHN 8693 (2, 55.3–57.4), Red Sea, Ehrenberg. Kenya: SAIAB 3106 (1, 85), Malindi, 2 Oct 1952, x-ray. SAIAB 3111 (1, 94), Shimoni, 1950, x-ray. SAIAB 3107 (2, 77–83) Shimoni, Nov 1952, x-ray. SAIAB 3112 (3, 15– 66), Shimoni, May 1951, x-ray. SAIAB 3109 (1, 76), Feb 1952, x-ray. Mozambique: SAIAB 3105 (1, 79), Pinda reef, 1950, x-ray. SAIAB 3114 (1, 49), Mocimboa de Praia, 3 Sep 1951, x-ray. SAIAB 3110 (1, 81), Inhambane, Sep 1948, x-ray. SAIAB 3108 (3, 59–82), Jun 1950, x-ray. Aldabra: SAIAB 3113 (1, 55), Nov 1954, x-ray. Apogon thurstoni Holotype BMNH 1889.8.17.2 (1, 55.5), India, Madras, Day. Amia uninotata Holotype USNM 70248 (1, 43.9), Philippines, Bisucay I., near Cuyo, Albatross, 9 Apr 1909, x-ray. Paratype USNM 339043 (1,40.0), Philippines, Palawan Is., Tara I., Mindoro Strait, Albatross, 15 Dec 1908, 3– 6 m., x-ray. Quinca mirifica Holotype WAM P5787.001 (1, 78.0), Australia, Western Australia, Cockatoo Island, Yampi Sound.

Apogonichthyoides nigripinnis species complex: The nigripinnis complex is characterized by: pale or light tan to almost brownish-black body with two prominent or diffuse dark bars, often with an ocellated spot as part of the anterior body bar; for adults without obvious body bars young or juveniles will have bars; darkish caudal peduncle bar or spot or may only be present in young and juveniles; head without light or dark snout striping, with brownish or blackish markings on the cheek and sometimes behind the eye; anterior part of first-dorsal fin pale to blackish or all of fin completely brownish-black; rounded soft dorsal without a continuous stripe, may have darkish base and pale-whitish distally; rounded anal fin without a stripe; emarginate to truncate pale to whitish caudal fin; pelvic fin spine often whitish; low number of welldeveloped gill rakers; tubular anterior nare.

Apogonichthyoides timorensis species complex: The timorensis complex is characterized by: brownish bodies with variable spots and/or darkish bars on the body without ocellus in anterior body bar; head lacking light or dark snout striping, with brownish or blackish markings on the cheek and behind the eye; blackish mark in first-dorsal fin; rounded soft dorsal with darkish stripe, stripe-like markings or darkish spot; rounded anal fin without a stripe; emarginate caudal fin with tiny brownish marks on fin rays, whitish or yellowish area on base of pelvic fins; low number of well-developed gill rakers; tubular anterior nare.

Figures

FIGURE 1. Apogonichthyoides uninotatus. A. Holotype of Amia uninotata, USNM 70248, 43.9 mm SL, by Violet Drandridge. Smithsonian Institution, NMNH, Division of Fishes. B. Postmortem buccal male, USNM 395671, 41 mm SL, BUS 03 - 07, Philippines, by J. T. Williams. C. Live, Coron I., Calamianes Is., Philippines, by G. R. Allen.