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Accepted by L. A. Rocha: 15 Nov. 2007; published: 29 Jan. 2008 1
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2008 · Magnolia Press
Zootaxa 1691: 1–52 (2008)
www.mapress.com/zootaxa/
Cardinalfishes of the genus Nectamia (Apogonidae, Perciformes) from the
Indo-Pacific region with descriptions of four new species
THOMAS H. FRASER
Mote Marine Laboratory, 1600 Ken Thompson Parkway, Sarasota, FL 34236-1096 USA.
E-mail: cardinalfish@comcast.net
Table of contents
Abstract ...............................................................................................................................................................................1
Introduction .........................................................................................................................................................................2
Methods and materials ........................................................................................................................................................3
Nectamia .............................................................................................................................................................................4
Key to the adult species of the Nectamia ............................................................................................................................7
Nectamia annularis (Rüppell, 1829) ............................................................................................................................8
Nectamia bandanensis (Bleeker, 1854) .....................................................................................................................13
Nectamia fusca (Quoy and Gaimard, 1824) ...............................................................................................................15
Nectamia ignitops new species ..................................................................................................................................23
Nectamia luxuria new species ....................................................................................................................................26
Nectamia savayensis (Günther, 1872) ........................................................................................................................32
Nectamia similis new species .....................................................................................................................................37
Nectamia viria new species ........................................................................................................................................41
Nectamia zebrinus (Fraser, Randall and Lachner, 1999) ...........................................................................................44
Color as a basis for species relationships ..........................................................................................................................46
Discussion .........................................................................................................................................................................48
Acknowledgments .............................................................................................................................................................50
Literature cited ..................................................................................................................................................................51
Abstract
The status of Nectamia is re-examined. General morphology including gill arches, cephalic lateralis pores, dorsal-fin ele-
ments, scale characteristics and results from recent contributions support recognition as a genus rather than a subgenus.
Nine species are removed from Ostorhinchus and placed in Nectamia. Neotypes are established for Apogon fuscus, the
type species of Nectamia and for N. bandanensis. Four undescribed species, Nectamia ignitops, N. luxuria, N. similis,
and N. viria, having been confused with N. bandanensis, N. fusca and N. savayensis. Two species, Nectamia annularis
and N. zebrinus are endemic to the Red Sea. Meristics or morphometrics, absent good color patterns will not conclu-
sively identify each species. A key to the species is provided. Four species are polymorphic with respect to number of gill
rakers. Gill-raker variation can be associated with geographic patterns: Red Sea, East Africa plus Mascarene Plateau,
China Sea, Indonesia plus Philippines, Palau to Wake I., and Southeast Indonesia-Coral Sea to Mangareva. Color pat-
terns are used in an analysis of phylogeny of the species. Five names are treated as synonyms: Apogon erdmani, a syn-
onym of Nectamia annularis and Apogon guamensis, Apogon nubilus, Apogon ocellatus, and Apogon spongicolus, as
synonyms of N. fusca.
Key words: cardinalfishes, new species, neotype, phylogeny, Apogon, Ostorhinchus, Pristiapogon, Pristicon, Zoramia,
Apogon annularis, Apogon bandanensis, Apogon erdmani, Apogon fuscus, Apogon guamensis, Apogon savayensis,
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Apogon nubilus, Apogon ocellatus, Apogon spongicolus, Apogon zebrinus, Nectamia, Nectamia annularis, Nectamia
bandanensis, Nectamia fusca, Nectamia ignitops, Nectamia luxuria, Nectamia savayensis, Nectamia similis, Nectamia
viria, Nectamia zebrinus
Introduction
Fraser (1972) recognized Nectamia Jordan, 1917 as a subgenus within Apogon Lacépède, 1802a with many
species whose relationships were poorly understood. Nectamia contained more species than any other subge-
nus. Gon (1987) fixed Ostorhinchus Lacépède, 1802b with a neotype and by this action, assumed priority over
Nectamia. Bergman (2004) noted some variation in pore patterns among the five species examined in
Ostorhinchus. Her cladistic analysis of the generic and subgeneric relationships of apogonids did not capture
any additional groupings within Ostorhinchus. Based on Bergman’s dissertation, Randall (2005) and Green-
field et al. (2005) recognized Ostorhinchus and Zoramia at the generic level. Mabuchi et al. (2006) examined
molecular phylogenetic information associated with color patterns for species of Apogon, sensu lato. Three
groupings were recovered among the 32 species examined in Ostorhinchus. Group II (barred pattern) can be
associated with an available name, Nectamia. Group III (striped pattern) can be associated with an available
name, Ostorhinchus. Group I species was recovered with another subgenus, Zoramia. Given the conflicting
hypotheses, the basis for recognizing genera or subgenera and their relationships in or out of Apogon, sensu
lato remains unstable. Bergman (2004) stated in her summary of conclusions that ‘...the weakly supported
phylogenies provide very little information upon which to make necessary taxonomic revisions’. What has
been clear for a long time is the need to examine a large number of species along with the identification of
phyletically useful characters. Part of Bergman’s dilemma was the sparse representation of species she exam-
ined in Ostorhinchus for cephalic lateralis characters.
Perhaps, some presently recognized subgenera of Apogon should be recognized at the generic level. Phyl-
ectic hypotheses by Bergman (2004) and Mabuchi et al. (2006) shows support for recognizing the following
monophylectic groups: Apogon (see Fraser and Randall, 2002), Pristiapogon (see Fraser and Lachner, 1985),
and Pristicon (see Randall and Fraser, 1999). Ostorhinchus, sensu lato, (Fraser, 1972; Bergman 2004; Mabu-
chi et al. 2006) by default, remains a polyphyletic grouping in need of more study. The divisions of Ostorhin-
chus suggested by Mabuchi et al. (2006) provide a rational basis for further testing of monophyly.
Examination of species in Zoramia and Ostorhinchus group I for morphological evidence of phyletic relation-
ships is an on going study in association with new species description.
The only review of the ‘bandanensis’ group of Apogon was done by E. A. Lachner in 1951. He recog-
nized four species. Prior to Lachner’s work, most ichthyological studies treated this complex as a polychromic
group of one or two species (Bleeker, 1874, 1871–1876; Jordan and Seale, 1906, Fowler and Bean, 1930;
Weber and de Beaufort, 1929; Schultz, 1943). Yet, much uncertainty remained even in Lachner’s discussion
of Apogon savayensis and Apogon bandanensis (1951, p.599–600). He correctly concluded that more study
with fresh material from the East Indies and the Philippine Islands and elsewhere was needed.
Five species were described between 1824 and 1872. Quoy and Gaimard (1824) described Apogon fuscus
from Guam: however no types existed according to Bauchot and Desoutter, (1987). The original description
of the caudal-peduncle mark was not completely diagnostic for species identification. Jordan and Seale (1906,
p244, fig. 34) and Jordan (1917) confused Apogon fuscus with another recently described species (see Green-
field, 2007), creating more uncertainty about the concept of this species. Rüppell (1829) described Apogon
annularis from the Red Sea. An endemic to the Red Sea, this species was occasionally confused with other
Indo-Pacific species. Valenciennes (1832) described Apogon guamensis based on a single specimen from
Guam. This wide-spread species was described several times. Bleeker (1854) described Apogon bandanensis
from two specimens collected at Banda Island. Confusion about the identity of Bleeker’s species was wide
spread in the literature and the name has been associated with described and undescribed species. Günther
Zootaxa 1691 © 2008 Magnolia Press · 3
status of Nectamia & 4 new species
described Apogon savayensis form Savai’i in 1872. Specimens of this species were confused most often with
two undescribed species. Four new species descriptions since Günther’s description have been determined to
be synonyms of older names (Fraser et al. 1999).
General usage of the ‘bandanensis’ group names has been chaotic because of a presumed lost type, two
type specimens mixed with non-type specimens, most types having almost completely faded color patterns,
some types without original illustrations, incomplete descriptions, and overlapping distribution patterns show-
ing all type localities with more than one species. Reliance by subsequent ichthyologists on poorly preserved
specimens or limited material created many misidentifications. While a historical chronicle of these five early
names and their synonyms might have some merit, the high level of uncertainty for specimen identification
from most past literature would require a herculean endeavor.
Species concepts for two of these Indo-West Pacific species (A. bandanensis and A. savayensis) were not
well understood in 1974 when I began a collaboration with E. A. Lachner, primarily the result of undescribed
species identified with these two names. Collecting and photographic materials by many ichthyologists during
the past 33 years have yielded material which helped to resolved some of the distinctive, yet subtle color pat-
terns. A recent review of three species occurring in the Red Sea was treated by Fraser et al. (1999). The Red
Sea endemics, Nectamia zebrinus and N. annularis, will not be redescribed in detail. Nine species are recog-
nized in this review. Several species have geographic variation in gill-raker counts indicating some degree of
polymorphism perhaps associated with partial isolation. Future studies may provide evidence for recognizing
additional species or subspecies.
Methods and materials
Methods for meristic data and measurements are given in Fraser (2005). In the dorsal spine formula, a spine
inside a parentheses (I) means this spine is hidden below the skin. Meristic data and proportions as % of stan-
dard length (SL) are given for the holotype, lectotype or neotype followed by the ranges in parenthesis. Senior
and junior synonymies are presented but reference lists associated with synonyms are not given because of the
inability to resolve identities solely from many publications. Some material examined, but not reported here,
was not allocated because of poor condition. Acronyms used to designate institutions and collections cited fol-
low Leviton et al. (1985) and Eschmeyer (1998).
Morphological characteristics were re-examined for Apogon fuscus, the type species of Nectamia and the
other species treated herein. Comparisons were made with the type species for Ostorhinchus, O. fleurieu, and
other related species to determine if supporting characteristics were present for recognizing Nectamia as
another lineage. Additional external characters were surveyed across all recognized subgenera to aid in devel-
oping a characterization of Nectamia. The species Mabuchi et al. (2006) recognized in the three grouping or
possible additional groups within Group II and III require a more complete survey along with additional gen-
era. References herein to other valid species referred to Apogon (Ostorhinchus) are used in the sense of Fraser
(1972) except where noted. Placement of these species may be uncertain as to groupings which may or may
not be allocated to a monophylectic Ostorhinchus. Specimens of Ostorhinchus fleurieu examined include
USNM 171272, 79.1 mm SL, cleared and stained, USNM 213216 and USNM 191704. One specimen of
Ostorhinchus aureus, USNM 212426 was examined. Twenty-four species other species in Ostorhinchus were
examined for head, body, pelvic, and lateral-line scales and second gill-arch characteristics.
The species are treated in alphabetic order. In the species accounts, material examined is listed by major
geographic areas and countries. Data referring to type specimens, including those pertaining to primary syn-
onymies, are identified and treated separately. Synonymies are based on examination of type specimens, orig-
inal description (or redescription of a type) and original illustrations. Collection localities from material
examined are plotted on maps. The symbols on the maps can represent one or more collections taken at differ-
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ent times and one or more specimens. Several maps include identified areas with averaged gill-raker counts.
Gill-raker counts were broken down into rudiments and rakers. Count variation presented in the geographic
tables include the sum of both elements. Oral incubation was checked by examining the mouth cavity of spec-
imens with buccal enlargement. The size of specimens with eggs in the mouth is listed under the remarks sec-
tion. All photographs were processed through Photoshop CS ver. 9.02 software. Other figures were obtained
from Wild camera lucida tracings, scanned on a flatbed scanner and finalized in Photoshop. Paup software
version 4.0b10 was used to develop hypothesized relationships derived from a suite of characters (Swafford,
2005). Output from Paup was processed in TreeViewX ver. 0.5.0 software.
Systematic accounts. Nectamia is distinguished from other species placed in Apogon, sensu lato, or more
recently from Ostorhinchus, sensu lato. Nine valid species are recognized based on color patterns, gill-raker
counts, and morphometrics. Nectamia, with its type species Apogon fuscus, is fixed by virtue of establishing a
neotype from the collection that Quoy and Gaimard made in Guam (see discussion under N. fusca). A neotype
is established for Apogon bandanensis from Bleeker’s putative, but mixed, type collection. Five nominal spe-
cies have been placed in synonymy, four in the wide spread Nectamia fusca and one in N. annularis (Fraser et
al. 1999). The number of new species is unusual for a well- collected, long recognized group — a function of
the previous confusing polychromic status accorded by earlier ichthyologists.
Nectamia Jordan, 1917
Figures 1–6, Tables 1–3
Type Species Apogon fuscus Quoy and Gaimard, 1824
Diagnosis. Species with IX dorsal spines as VII(I)–I, the eighth of nine dorsal spines reduced to a tiny
nubbin hidden under the skin supported by a free sixth distal radial (Fig. 1), dorsal spines slender, with slight
thickening of the third spine; 9 soft dorsal rays; 8 soft anal rays; 13 pectoral-fin rays; 23–30 total rudiments
and rakers on the first gill arch, 22–29 well-developed gill rakers, 16–18 shorter gill rakers present on the sec-
ond ceratobranchial and hypobranchial (Fig. 2); three supraneurals; two supernumerary spines on the first dor-
sal pterygiophore; one supernumerary spine on the first anal pterygiophore; one pair of slender uroneurals,
three epurals, free parhypural, five free hypurals; procurrent caudal rays segmented; forked caudal fin, all
head and body scales ctenoid, scales ctenoid or cycloid on base of pectoral fin, two large median pelvic scales,
the posterior scale cycloid in some species, no developed accessory pelvic scale (Fig 3), pored lateral-line
scales with one or two distal openings above and below main lateralis canal (Fig. 4); cephalic lateralis pores
numerous (Fig. 5), mostly as minipores, a terminal lachrymal pore, a pair of ventral lachrymal pores, an ante-
rior dentary and mental pores, terminal end of supraorbital canal with a large pore near anterior nare, lateral
margin of supraorbital canal near posterior nare small, lateral margin of supraorbital canal midway on interor-
bit with small pores, simple postorbital canal projections with minipores, anterior margin of supratemporal
canal smooth, posterior supratemporal canal margin with complex projections and minipores, lateral margin
of mandibular canal with many minipores, a small posterior articular pore; stomach and intestine black with
pale peritoneum; dark peduncular spot or bar, dark cheek mark between angle of preopercle and eye, lachry-
mal and anterior infraorbitals with yellowish mark, no stripes on head, body or vertical fins, pale bars and dark
saddles variable present or absent on body, dorsal and ventral caudal-fin edges pale to variously colored (Fig.
6, Tab.1).
Remarks. External color patterns are important distinguishing characters in this genus, particularly the
caudal-peduncle marks, dorsal saddles, cheek marks, development of pale bars on the side of the body, and
development of colored edges to the caudal fin. Historical and recent methods of handling postmortem speci-
mens prior to fixing and during preservation techniques influence overall preserved markings: kinds and
strength of alcohol, formalin in freshwater or saltwater, collection methods and the use of ice or cold water
Zootaxa 1691 © 2008 Magnolia Press · 5
status of Nectamia & 4 new species
prior to photography. The yellowish mark on the lachrymal and anterior infraorbitals was common to all spe-
cies with good underwater photographs. Such a lachrymal mark has not been described for other species com-
plexes in Ostorhinchus. Thirteen pectoral-fin rays are present in all species, an uncommon number among
species in Ostorhinchus, sensu lato. A pelvic axillary scale is absent, however, a longer than wide small rudi-
mentary cycloid scale was hidden by the larger ctenoid scale in some individuals.
FIGURE 1. Mid-dorsal fin bones and spine associated with the sixth proximal-medial element. (A) Nectamia fusca,
USNM 112348, 62.3 mm SL, connection between the first and second dorsal-fin elements with a free sixth distal radial
and tiny eighth dorsal spine. (B) Ostorhinchus fleurieu, USNM 171272, 79.1 mm SL, connection between the first and
second dorsal-fin elements with a fused distal radial to the seventh proximal-medial element and absence of an eighth
dorsal spine. Scale = 1.0 mm.
FIGURE 2. Gill rakers and rudiments on second arch. Ostorhinchus fleurieu, USNM 171272, 79.1 mm SL, (A) hypo-
branchial, dorsal view, (B) ceratobranchial, lateral view. Nectamia fusca, USNM 112348, 62.3 mm SL (C) hypobran-
chial, dorsal view, (D) ceratobranchial, lateral view. Scale = 1.0 mm.
A broader survey of species variation across Ostorhinchus, sensu lato, for patterns of the cephalic lateralis
pores, fused sixth dorsal radial (Fig. 1B) fewer developed gill rakers on the first and/or second arches (Fig. 2A
and B), complex lateral-line scale pores (Fig.3D), pelvic base scales and a pelvic axillary scale (Fig. 4B) will
determine usefulness in developing phyletic relationships. Here, simple pored lateral-scales, absence of a pel-
vic axillary scale and the absence of stripes on the head and body are enough to distinguish Nectamia in
Mabuchi et al. (2006) group II from Ostorhinchus in their group III. A broad survey may show that a fused
versus free sixth distal dorsal radial may be useful.
Nectamia is a well-resolved clade. Color pattern similarities among apogonid genera and within species
groups suggest an evolutionary basis that will be more widespread than Mabuchi et al. (2006) demonstrated
for some of the groupings in Apogon, sensu lato, including those species in their Ostorhinchus group II. Two
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other names are available for other branches of species placed in Group II: Jaydia as proposed by Smith
(1961) and used by Gon (1996), but see Fraser (2000); and Apogonichthyoides proposed by Smith (1949).
Nectamia is narrowly construed here: however, barred patterns and/or absence of head stripes may be enough
to include these other lineages within Nectamia which would broaden the diagnosis and eliminate some char-
acters used here to more strictly define Nectamia. A key to aid in the identification of species from preserved
material follows.
FIGURE 3. Semi-diagrammatic drawings of the fourth lateral line scales from left side of body showing variation in the
number of external peripheral pores. A. Nectamia fusca scale removed from body, USNM 139811, 60.2 mm SL. B.
Nectamia viria, holotype USNM 319149, 57.4 mm SL, scale on body. C. Nectamia similus, holotype USNM 213123,
44.0 mm SL, scale on body. D. Ostorhinchus fleurieu, USNM 213216, 63 mm SL, scale on body. E. Nectamia luxuria,
holotype, USNM 209644, 59.1 mm SL scale on body. F. Nectamia ignitops, holotype BPBM 21897, 68.1 mm SL, scale
on body. Scale = 1.0 mm.
FIGURE 4. Comparison of the scales at the base of the pelvic spine for A. Nectamia fusca without an axillary scale,
USNM 139811, 60.2 mm SL and B. Ostorhinchus fleurieu with an axillary scale, USNM 191794, 42.3 mm SL.
Zootaxa 1691 © 2008 Magnolia Press · 7
status of Nectamia & 4 new species
FIGURE 5. Semi-diagrammatic pattern of cephalic lateralis pores for Nectamia fusca, USNM 139811, 60.2 mm SL,
Guam. A. Dorsal view. B. Lateral view. C. Ventral view. d. last dentary/anticular pore. n. anterior and posterior nares. e.
posttemporal.
Key to the adult species of the Nectamia
1. Oblique cheek mark expanded near eye; basicaudal spot, saddle or bar; body depth generally greater than
42% of standard length................................................................................................................................ 2
- Oblique cheek mark narrow throughout length; basicaudal spot central and diffuse in adults; body depth
generally less than 42% of standard length. Indo-Central Pacific.................................................................
................................................................................................ Nectamia fusca (Quoy and Gaimard, 1824).
2. Lobes of the caudal fin with dark bands near upper and lower edges.........................................................3
- Caudal fin dusky, no dark bands in lobes near upper and lower edges .......................................................6
3. Well developed light bars on side of body, may have narrow or broad dark areas between the light bars. 4
- No well developed light bars, occasionally a few faint light bars. Western Indian - Central Pacific............
.......................................................................................................... Nectamia savayensis (Günther, 1872)
4. Light bar present on the opercle flap, a few narrow light bars on body, dorsal saddles under the first and
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second dorsal fins. West Pacific ................................................................................ Nectamia similis n.sp.
- No bar on the opercle flap, no dorsal saddle under the first and second dorsal fins ................................... 5
5. Pale bars with wide darker area on body, black caudal-peduncle saddle extends below lateral line but not a
complete uniform band. Red Sea-Gulf of Aden.. Nectamia zebrinus (Fraser, Randall and Lachner, 1999).
- Caudal-peduncle mark above lateral-line; pale bars with narrow dark bars on body. Indian-Central Pacific
.................................................................................................................................. Nectamia luxuria n.sp.
6. Dark caudal-peduncle band extend below lateral-line scales; dorsal saddles present or faint................... 7
- No pale bars; no dorsal saddles; broad cheek mark. Red Sea............ Nectamia annularis (Rüppell, 1829).
7. Two dorsal saddles, no pale bars on body................................................................................................... 8
- Dorsal saddle present under second dorsal fin; a few faint pale bars sometimes present. West Pacific .......
...................................................................................................................................... Nectamia viria n.sp.
8. Total gill rakers and rudiments usually 26–29. West Pacific..........Nectamia bandanensis (Bleeker, 1854).
- Total gill rakers and rudiments usually 24–27. South China Sea............................ Nectamia ignitops n.sp.
Nectamia annularis (Rüppell, 1829)
Figures 6E, 7, 18 and Tables 1–3
Synonyms: Apogon erdmani Lachner, 1951. Red Sea, Saudi Arabia, Jidda.
Type material. Lectotype Apogon annularis SMF 1774 60.9 mm SL; Red Sea, El Tur, Sinai coast, Egypt,
Gulf of Suez.; E. Rüppell, 1827. Paralectotypes SMF 4679-81 (3, 37–51); same date as SMF 1774. Holotype
Apogon erdmani USNM 147518 59.5 mm SL; Saudi Arabia, Jeddah, Sams Pier, 2 July 1948, D.S. Erdman
Coll; x-rayed. Paratypes USNM 112040 (7, 43–59); same data as holotype.
TABLE 1. Geographic variation in gill-raker and rudiments counts for Nectamia annularis.
* = lectotype
Additional Material. Red Sea: Egypt: Gulf of Aqaba: USNM 212429; (24, 39–53); Bay at El Himera.
BPBM 21513; (3, 34–45); color; El Himera; 12m.; 25 Apr. 1977. SMF 1076; (1, 73); 1872. SMF 4682; (1,
49); Al Ghardaqa, 27°14'N 33°50'E; 27 Jul 1957. USNM 212430; (16, 36–59); N. of Râs Burqa. USNM
212432; (5, 40–52); Marsa Muqabila. USNM 212438; (8, 42–50); One mi. N. of Râs Burqa. USNM 212441;
(1, 40); Between Marset Mahashel Ala & Marset Abu Samra. USNM 212443; (3, 14–36); Bay between marsa
Mokrakh and El Himera. Strait of Gûbâl: USNM 212428 (61, 45–59); Râs Muhammad. USNM 212439; (2,
31–50). USNM 212442; (21, 25–52). USNM 212444; (1, 26). Gulf of Suez: USNM 212436; (9, 26–30).
N=52 Upper Arch Lower Arch Total
5678x 19 20 21 22 x 24 25 26 27 28 29 x
Red Sea
Aqaba 74 7.4 3 7 1 20.8 2 5 4 28.2
El Tur 1 1 9* 1 6.8 1 6 5* 20.3 1 1 4 6* 27.2
Gulf of Suez 1 1 20.0 1
Isola Delemme 1 7 2 7.1 1 9 20.6 1 8 1 27.0
Jidda 12 2 7.1 1 9 4 19.9 1 7 5 1 27.4
Khor Obhour 2 7.0 1 1 20.5 1 1 27.5
Gold Mohur Bay 2 7.0 1 1 19.5 1 1 26.5
Total 1 2 40 9 7.1 4 30 17 1 20.3 1 0 4 24 18 5 27.4
Zootaxa 1691 © 2008 Magnolia Press · 9
status of Nectamia & 4 new species
USNM 212437; (30, 32–55). USNM 212440; (3, 28–57); El-Tur, Sinai Peninsula. USNM 212446; (72, 24–
58). Jordan: SMF 16161; (2, 32–32); Aqaba; 14 Nov 1979. SMF 16162; (1, 30); Aqaba; 10 Nov 1979; 10 m.
SMF 16163; (1, 51); Aqaba; 23 Nov 1979; 10 m. Saudi Arabia: ANSP 163226; (2, 52–53); Khor Obhour.
RMNH 12892; (2, 61–66); Jeddah. BPBM 30386; (4, 14–18); 29km. S. of Yanbu. Sudan: BPBM 20373; (3,
34–45); Suakin Harbor. SMF 8352; (1, 20); Suakin, 19°07'N 37°20'E; Sep 1964. Eritrea: USNM 212431;
(29, 27–55); North end of Isola Delemme, off Massawa. USNM 212433; (47, 24–53); Massawa. USNM
212434; (7, 17–29); Difnein I., South shore. USNM 212435; (2, 31–35); Melita Bay. USNM 212445; (1, 28).
Yemen: Red Sea: BPBM 35701; (4, 18–22); Jaz ’ir as Zubayr, 15°6.8' N., 42°35.9' E. Gulf of Aden: USNM
212447; (20, 20–62); Gold Mohur Bay.
This species is not treated in detail except for comparisons with other species, additional specimens and
new information under the Description. See Fraser et al. (1999) for additional information.
Diagnosis. A Nectamia without bars on body, circumpeduncular basicaudal mark in juveniles and adults,
caudal fin with dark margins, subocular mark broad, triangular (Tab. 2); 13 pectoral-fin rays; total gill rakers
usually 26–29 (Tab. 1); body depth 41–50%, caudal-peduncle depth 15–20%, second anal-spine length 18–
22%, and pectoral-fin length 26–31% of standard length (Tab 3).
FIGURE 6. Underwater photographs of eight species of Nectamia. A. N. similis, new species, Calamian Is., Philippines,
G. R. Allen. B. N. luxuria, new species, Andaman Sea, Thailand, U. Satapoomin. C. N. viria, new species, Raj Ampat,
Indonesia, R. F. Myers. D. N. zebrinus, Red Sea, Egypt. R. F. Myers. E. N. annularis, Red Sea, Sudan, J. E. Randall. F. N.
savayensis, Solomon Sea, Bougainville I., J. E. Randall G. N. bandanensis, Ogasawara Is., Japan, J. E. Randall. H. N.
fusca, Red Sea, R. H. Kuiter.
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TABLE 2. Preserved color pattern characteristics of the adult species of Nectamia.
Additional Description. Second gill arch with well developed rakers, 4+18; scales ctenoid on cheek, sub-
opercle, opercle, nape, isthmus, two large scales on base of pelvic fin, rudimentary axillary scale variable
present cycloid to few ctenii, cycloid on base of pectoral fin and ctenoid behind pectoral fin, pored lateral-line
scales complete, fourth scale with one pore above and one below main canal.
Preserved color pattern. Adults (Tab. 2): wide, triangular subocular cheek mark behind upper jaw; first
dorsal fin dark membranes between spines 1–4; edges of caudal lobes dark; caudal peduncle with a complete
dark wide bar, connected dorsally and ventrally, body uniform without pale bars; no dark saddle under the first
or second dorsal fin; stomach and intestine black. Juveniles: similar to the adults.
Life colors. Kuiter and Kozawa (1999, p.34, A–C) provided three of underwater photographs of this spe-
cies. Gon and Randall (2003, Pl.1A) published a photograph of a freshly collected specimen. An underwater
photograph (Fig. 6E) shows the yellowish mark on the lachrymal and anterior infraorbitals, pale body bars,
dark edges of dorsal fins, cheek mark and caudal-peduncle marks.
Distribution. Known only from the Red Sea and the Gulf of Aden (Fig. 7).
annularis viria bandanensis similis fusca savayensis zebrinus luxuria ignitops
pale barring on
side of body none none to
few faint none few none to
few faint none to
few faint many many none
pale bar on none none none one bar none none none none none
opercle
lobe edges of no dark no dark no dark dark no dark dark dark dark no dark
caudal fin stripe stripe stripe stripe stripe stripe stripe stripe stripe
caudal-peduncle
mark
young complete complete complete complete spot partial complete partial unknown,
band band band band band band band (complete
band?)
adult complete complete incomplete to incomplete diffuse incomplete incomplete incomplete complete
band band complete band band spot band band band band
cheek mark wide, narrow, narrow, narrow, narrow wide, wide, wide, narrow
triangular triangular triangular triangular line triangular triangular triangular
dark dorsal
saddles none under 2nd
dorsal fin under 1st and
2nd dorsal fin under 1st and
2nd dorsal fin none none or under
2nd dorsal fin none none under 1st
and
2nd dorsal
fin
roof of pale pale blackish numerous pale pale pale pale blackish
mouth melanophores
upper arch absent absent present present absent absent absent absent present
melanophores
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status of Nectamia & 4 new species
FIGURE 7. Distribution of the material examined for Nectamia annularis and N. viria with regional total gill raker
averages.
Remarks. See species comparison under remarks for Nectamia zebrinus. Günther (1859) used the com-
bination Apogon annularis var. roseipinnis. Klausewitz (1959) suggested that Apogon annularis is a subspe-
cies of Apogon aureus. These subspecies concepts have not been supported by any subsequent work.
Klausewitz (1959) and Smith (1961) concluded that Apogon erdmani is a synonym of Apogon annularis and
Fraser et al. (1999) came to the same conclusion.
Two males in USNM 212429 exhibited buccal enlargement but no eggs were found in their mouths. This
species has been collected at the same stations with Nectamia zebrinus (BPBM 19741, 30388, USMN
212423) in the Red Sea and Gulf of Aden (USMN 212421). The largest of 400 specimens was 66 mm SL.
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TABLE 3. Proportions as a percent of standard length for species of Nectamia. The proportions for primary types are followed
by parentheses for other types and additional material. The hyphen is a missing (unavailable) measurement from a specimen.
Body Proportions annularis* viria bandanensis† similis fusca‡ savayensis zebrinus luxuria ignitops
percent of standard length
greatest body depth 45.4(41-50) 44.2(44-47) 40.4(38-46) 44.8(43-45) -(37-42) 41.7(40-45) 44.8(40-47) 43.0(39-44) 46.1(44-47)
head length 41.9(41-46) 43.2(43-45) 39.3(43-48) 44.5(42-46) -(40-44) 42.4(41-45) 45.8(43-52) 42.1(42-45) 44.2(43-44)
Eye diameter 15.9(16-18) 16.5(16-18) 14.9(16-20) 17.0(16-19) -(14-18) 16.1(13-17) 16.9(14-18) 16.4(15-16) 16.1(16-17)
snout length 10.2(8-10) 8.5(8-10) 8.7(9-11) 9.1(9-10) -(8-9) 8.5(8-10) 8.5(8-11) 8.5(7-9) 9.5(8-10)
bony interorbital width 11.1(10-11) 11.1(10-12) 10.0(10-13) -(10-13) -(10-12) 11.6(9-12) 12.1(10-14) 11.0(10-12) 11.0(10-12)
upper jaw length 21.6(21-22) 20.2(19-21) 19.2(22-24) 21.8(21-22) -(20-22) 20.7(18-21) 21.8(20-25) 19.5(19-23) 21.9(22-23)
caudal-peduncle depth 19.4(15-20) 18.8(17-19) 18.1(17-19) 19.5(18-21) -(15-18) 20.0(18-21) 19.6(18-22) 18.3(16-20) 19.5(18-20)
caudal-peduncle length 22.9(15-23) 22.3(21-24) 22.5(21-26) 18.2(18-22) -(22-26) 22.1(21-25) 21.5(18-24) 23.5(21-24) 21.7(19-24)
1st dorsal-fin spine length 3.0(2.5-5.3) 3.0(3-4) 3.4(2-4) 2.3(2-3) -(2.1-4.2) -(1.3-4.1) 2.5(2-4) 3.2(2-4) 3.2(3-4)
2nd dorsal-fin spine length -(9-12) 9.2(9-11) 7.9(9-12) 10.2(8-10) -(8-11) -(8-11) 11.2(10-14) 9.8(8-11) 10.1(9-12)
3rd dorsal-fin spine length 20.8(17-21) 18.3(18-21) 18.8(17-23) 18.2(17-19) -(17-22) -(16-23) 21.1(18-23) 17.3(17-22) 20.2(19-21)
4th dorsal-fin spine length 21.1(18-21) 18.3(17-21) 18.0(17-21) 20.0(17-20) -(16-19) -(17-21) 20.3(19-24) (17-20) 21.4(21)
2nd dorsal-fin spine -(17-20) 16.9(16-18) 17.7(17-21) 18.2(17-19) -(15-19) -(14-21) 17.6(17-21) 17.3(16-21) 19.2(19)
1st anal-fin spine length 5.4(3.4-5.4) 3.7(4) 4.1(3-6) 4.3(4) -(3-5) 2.9(3-4) 3.7(2-7) 3.4(3-5) 5.1(5)
2nd anal-fin spine length 20.8(18-22) 16.2(16-18) 15.3(18-22) 19.5(19-21) -(15-17) -(14-20) 19.6(19-23) 17.8(17-21) 19.2(19-21)
pectoral-fin length -(26-31) 26.1(26-29) 23.0(25-31) 26.8(26-30) -(24-26) -(25-29) -(26-32) 27.4(26-29) 28.6(25-29)
pelvic-fin length 25.8(25-26) 24.4(24-26) 23.4(24-27) 26.6(25-27) -(21-26) 17.6(24-26) -(24-31) 26.1(24-27) 25.3(25-28)
*SMF 1774, 59.7 mm SL lectotype.
†RMNH 5584, 71.2 mm SL, neotype.
‡MNHN 8682, 52.0 mm SL, neotype.
Zootaxa 1691 © 2008 Magnolia Press · 13
status of Nectamia & 4 new species
Nectamia bandanensis (Bleeker,1854)
Figures 6G, 8, 18 and Tables 2–4
Synonyms: misspelled as Apogon batjanensis by Weber 1895.
Type material. Neotype Apogon bandanensis RMNH 5584, 71.2 mm SL, 90+ total length, Indonesia, Banda
Neira, one of 29 specimens (17–80 mm SL, 24-103 TL).
Additional material. Indonesia: BMNH 1880.4.21.83–86; (4, 51–71); Banda Neira. Papua New
Guinea: Coral Sea: BPBM 32437; (2, 66–71); off Bootless Inlet, S. of Motupore I., caves; 13–15m; 28 Oct
1987; x-ray. Fiji: CAS 220918; (2, 55–60); Charybdis Reef, 17°13.138'S, 178°02.851'E; G03-78; 30 May
2003; 14–18 m. CAS 223740; (1,69); Pascoe Reefs, W of Yadua I., 16°50.188'S, 178°13.665'E; G02-136; 4
Apr 2002. CAS 223742; (3, 47–60); Patch reef, 17°13.138'S, 178°02.851'E; G03-82; 30 May 2003; 12–18 m.
CAS 223743; (2, 59–60); NE of Yaqaga I., 16°30.717'S, 178°38.730'E; G02-104; 25Mar2002; 9–12 m. CAS
223754; (4, 52–59); outside barrier reef, 16°27.333'S,179°40.250'W; G03-55; 22 May 2003; 12–17 m. CAS
223755; (1, 60); Budd Reef, 16°30.444'S,179°40.571'W; G03-53; 22 May 2003; 20–24 m. Rotuma: USNM
287945; (4, 35–53); Hatan I.; VGS 86-14; 19 May 1987; x-ray. USNM 287966; (1, 43); VGS 86-16; 20 May
1986; 29–32 m; x-ray. Samoa: American Samoa: BPBM 17522; (3, 31–53); Tutuila; 9 May 1974; color
photo; x-ray. Guam: ANSP 152918; (3, 42–44); 20–30m; 27 June 1968. BPBM 17768; (2, 24–41); 27 June
1968; x-ray. Japan: Ryukyu Islands: BPBM 22308; (4, 34–75); Okinawa, Sesoko I.; 13 Sep. 1977; color
photo; x-ray. Marshall Islands: BPBM 28971; (1, 51) and BPBM 28974; (2, 51–53); Enewetak Atoll; 23m;
24–25 Sep. 1982; x-ray.
Diagnosis. A Nectamia with two dark saddles on body, one below the first dorsal fin, the second below
the second dorsal fin, basicaudal saddle extends below lateral-line scales in adults, no pale bars on side of
body, caudal fin without dark margins, subocular mark, thin and triangular (Tab. 2); 13 pectoral-fin rays; total
gill rakers usually 26–28 (Tab. 4); body depth 38–46%, caudal-peduncle length 21–26%, second anal-spine
length 18–22%, pectoral-fin length 25–31% of standard length (Tab. 3).
FIGURE 8. Distribution of the material examined for Nectamia bandanensis and N. ignitops .
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Description. Range of proportions (as percentage of standard length) in Table 3 with neotype first and
additional material in parentheses.
Dorsal fin VII(I)–I,9; anal fin II,8; pectoral-fin-rays 13-13; pelvic fin I,5; principal caudal rays 9+8; pored
lateral-line scales 25 (24); transverse scale rows above lateral line 2; transverse scale rows below lateral line
6; 4 median predorsal scales; circumpeduncular scale rows 12 as 5+2+5; total gill rakers 25 (25–29), well
developed 22 (22–27), upper arch 3+4 (1-3+5-7), lower arch 18+0 (18-21+0-1), second arch with shorter gill
rakers 3+18.
Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and
vomer; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10+14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural.
Three supraneurals, two supernumerary spines on first dorsal pterygiophore. Basisphenoid present. Supra-
maxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal mar-
gins. Infraorbital shelf present on third bone.
Scales ctenoid on cheek, subopercle, opercle, nape, isthmus, cycloid on base of pectoral fin and ctenoid
behind pectoral fin and rest of body. Two large ctenoid scales on base of pelvic fin, no axillary scale. Pored
lateral-line scales complete, fourth scale with one pore extending up to near overlapping scale edge and one
below main canal.
Life colors. See Figure 6G. Kuiter and Kozawa (1999) provide a number of underwater photographs (p.
33, A–E) identified as Apogon bandanensis. Their figure D of a Sulawesi specimen is the only one that I can
identify as this species. Their photographs of species 17, (p33, B and C) and species 19 (p.34, A and B) may
also represent this species. Randall’s photograph of a fresh specimen (BPBM 22308, 48 mm SL) has a red iris
and three blackish body bars, one from the base of the first dorsal fin reaching to behind the pectoral-fin base,
the second from the posterior base of the second dorsal reaching almost to the posterior portion of the anal fin,
the third a circumpeduncular bar, body otherwise silvery. A blackish membrane (distal two thirds) between
the third and fourth first dorsal spines, membranes dusky (darker distal two thirds) between the fourth and
seventh spines and dusky between the second and third spines. Second dorsal fin, anal fin and caudal fins uni-
formly pale. Darkish subocular cheek mark present.
TABLE 4. Geographic variation in gill-raker and rudiments counts for Nectamia bandanensis.
* = neotype
Preserved color pattern. Adults (Tab.2): narrow, triangular subocular cheek mark behind upper jaw; first
dorsal fin with dark membrane between spines 1–4; edges of caudal lobes plain; caudal peduncle with a nearly
complete dark wide bar, connected dorsally and almost connected ventrally; dark saddles under the first and
N=30 Upper Arch Lower Arch Total
6789x18 19 20 21 x25 26 27 28 29 x
Okinawa 4 8.0 1 2 1 20.0 1 2 1 28.0
Banda I. 1* 1* 1*
New Guinea 2 7.0 2 20.0 2 27.0
Guam 4 1 7.2 2 3 20.6 2 2 1 27.8
Fiji / Rotuma 1 9 3 7.2 5 5 3 19.8 5 5 1 2 27.0
Enewetak 1 1 8.5 2 20.0 1 1 28.5
Tutuila 1 2 7.7 3 20.0 1 2 27.7
Total 1 17 11 1 7.4 1 6 16 7 20.0 1 5 11 8 5 27.4
Zootaxa 1691 © 2008 Magnolia Press · 15
status of Nectamia & 4 new species
second dorsal fins; stomach and intestine black; roof of mouth dark with melanophores extending down onto
upper part of lower gill arch. Juveniles: similar to the adults, except with a complete, dark caudal-peduncle
band.
Distribution. Known only from the West Pacific, extending into the Central Pacific to the Marshall
Islands and Samoa (Fig 8), with a tendency to be found in deeper water (13–27m). This species has been col-
lected from Halimeda algae beds or rubble and dense corals.
Remarks. Bleeker mentions two specimens (75 and 94mm total length) in the original description. Ernest
A. Lachner (museum notes) and I independently examined RMNH 5584, the probable type series, which con-
tained 29 specimens, 18–81mm SL. Bleeker’s auction catalogue listed 24 specimens as do his 1874 and 1876
papers. No specimen in RMNH 5584 was close to the 75mm total length of one syntype and is considered
missing. One specimen, 81 mm SL was 90+mm total length was the closest to the 94 mm specimen and could
be considered to be the larger syntype. It is completely faded, lacking all of the distinguishing color pattern.
The British Museum has three specimens, 51–53 mm SL (all with broken caudal fins, total lengths unknown
54+, 61+ and 65+) and one specimen 70 mm SL (85+ TL) in BMNH 1880.4.21.83–86 listed as syntypes (also
in Eschmeyer, 1998). All the BMNH specimens are considered to be non-type material because none agrees
with the syntypes’ total lengths. All specimens belong in the species group but lack species specific color pat-
terns because of fading with time.
Additional Bleeker material is present in both museums beyond what Bleeker originally reported or in the
auction catalog (see Lamme, 1973). Because of the additional material, missing one syntype, and inability to
make a completely positive identification of a remaining syntype, designation of a lectotype is not possible.
The basis for Bleeker’s name is from data in the original description and his illustrations in the Atlas (1875–
76, percoid Pl. 67, Fig. 2). The iris of the eye is reddish in this figure and Bleeker (1874 and 1871–75) men-
tions this trait (yellow or red) but not in the original description: “... aureo fuscescente plus minusve arenato;
fasciis 3 latis transversis diffusis violascentibus, anteriore sub pinna dorsali spinosa, 2a sub pinna dorsali
radiosa, 3a cauda basi pinnae caudalis approximata; pinna dorsali radiosa fusca; pinnis ceteris roseis, caudali
leviter violascente.” The species treated here as Nectamia bandanensis requires the fewest assumptions and
has the best fit to Bleeker’s original concept and subsequent figure. The 90+mm total length specimen is more
likely then any other extant specimens to be one of the syntypes, but because the collection was contaminated
with non-type material a neotype is designated.
This species has been mis-identified from many locations by virtually all ichthyological reports except for
a few of the most recent publications. Lachner (1951) treated this species but provided a photograph (Plate
18,B) of a new species, Nectamia similis. His study material was from the Albatross Expeditions to the Philip-
pines and Indonesia and consisted of Nectamia luxuria (13 lots), N. savayensis (2 lot, both questionably
referred, p.599), N. similis (7 lots) and N. viria (1 lot, questionably referred, p.599). No specimens of
Nectamia bandanensis were clearly present. This species has been taken with Nectamia savayensis in the
Marshall Islands (BPBM 28971). The largest of 55 specimens was 80 mm SL.
Nectamia fusca (Quoy and Gaimard, 1824)
Figures 1A, 2C&D, 3A, 4A, 5, 6H, 9, 18 and Tables 2–3, 5
Synonyms: Apogon fuscus Quoy and Gaimard, 1824, Western Pacific, Guam. Apogon guamensis Valenciennes, 1832,
Western Pacific, Guam. Apogon nubilus Garman, 1903 Western Pacific, Fiji, Suva reef. Apogon ocellatus Fourman-
oir and Crosnier, 1964 Indian Ocean, Madagascar I., Nosy-Bè, Ambatoloaka. Ostorhynchus spongicolus Smith,
1965. Red Sea, Ethiopia.
Type material. Neotype Apogon fuscus MNHN 8682; 52.5 mm SL, 69.9 mm total length; Guam L. de Frey-
cinet, 1817–1820 (one of three spec., 52.0–58.0, largest with tag). Syntypes Apogon guamensis MNHN 8767
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16 · Zootaxa 1691 © 2008 Magnolia Press
4(22.2–56.0); Guam; Quoy and Gaimard; 1826–1829. Holotype Apogon nubilus MCZ 28315 (53.6); Fiji,
Suva reef; 12 Dec. 1897. Syntypes Apogon ocellatus MNHN 1973–40 2(20.1–20.3); Madagascar I., Nosy-Bè,
Ambatoloaka. Holotype Ostorhynchus spongicolus SAIAB 354 (17.0); Red Sea; Ethiopia; x-ray. Paratypes
SAIAB 441; 2(15.7); Red Sea; Ethiopia; x-ray. SAIAB 774; (22,1);Kenya, Shimoni; Nov.1952; x-ray.
Additional material. Red Sea: Israel: Gulf of Aqaba. SAIAB 3278 (1, 57); USNM 212876; (5, 64–75);
USNM 212875; (5, 47–63); USNM 212874; (3, 52–64); USNM 212878; (7, 59–71); USNM 191707; (4, 21–
28); Eilat; USNM 212873; (1, 67); USNM 212877; (12, 16–66) Muqabila; Egypt: SMF 19891; Sinai; (7, 15–
28); Oct 1973. USNM 212883; (10, 39–69); Strait of Jubal, 0–5m. USMN 212884; (1, 31);Strait of Jubal, S.
end of Sinai Peninsula. BPBM 18346; (1, 33); Râs Muhammed; 19 Sep. 1974. USNM 213617; (57, 32–71);
27°18'50"N, 33°47'35"E; 0–5m. USNM 212879; (6, 23–66). USNM 212880; (46, 33–68). USMN 212884; (1,
31); Harghada; USNM 212885; (1, 32); Ghardaqa; USNM 147519; (28, 53–68); NW edge Shaib Al Zanadir
Reef. Saudi Arabia: Jidda. USNM 191658; (3, 48–55); USNM 147523; (15, 12–66); USNM 112046; (12,
33–65); USNM 212886; (3, 40–65); USNM 163808; (1, 11); USNM 147524; (7, 37–60). Ethiopia: Sharm
Ubhar: USNM 212881; (9, 49–60); N. end of Isola Delemme. USNM 212882; (1, 49); naval base, Massawa.
Indian Ocean: Kenya: SAIAB 3280; (1, 48). SAIAB 3269; (1, 68); Shimoni. SAIAB 3277 (3, 23–31); Mom-
basa. BPBM 27212; (1, 69); NE of Mombasa; 15 March 1979. Zanzibar: SAIAB 3271 (2, 48–53); x-ray.
SAIAB 3273 (1, 37); x-ray. SAIAB 3281 (3, 62–66); x-ray. Ta nza ni a: SMF 4762; (1, 31); Mikindani Bay;
1909/10. Mozambique: SAIAB 3272 (1, 33); Cabo Delgado; x-ray. SAIAB 3274 (1, 42) and 3276 (1, 33);
Pinda Reef; x-ray. SAIAB 3275 (1, 47); Mozambique I.;x-ray. Madagascar: USNM 212886; (7, 10–22);
Nossi-Bé. USNM 212889; (2, 16–19); Nossi Be;18 Jan 1964. USNM 212888; (5, 10–23); Nossi Be; 3 Jan
1964. Comoro Is.: USNM 212887; (5, 12–21); Pamanzi I. Maldive Is.: CAS 35379; (170, 24–71); Male
Atoll, Hulele I.; 8 Nov 1964. CAS 56679; (2, 43–51); Male Atoll, Hulele I.; 8 Nov 1964; Te Vega 64–5.
FMNH 75710; (11, 27–65); Addu Atoll; 10 May 1964; LPW-40. FMNH 75659 and 75664; (51, 9–63); Fadif-
folu Atoll; 24 Mar 1964; LPW-27. FMNH 75654; (9, 17–65); S. Male Atoll; 18 Mar 1964; LPW-24. FMNH
75705; (8, 30–36); Addu Atoll; 8 May 1964. SMF 4751; (19, 30–53); Rasdu Atoll, W. Weligandu I. 13 Mar
1958. SMF 4763; (1, 43); Addu Atoll, Hittadu I.; 6 Jan 1958. India: FMNH 75599; (5, 20–73) Gulf of Man-
nar; 30 Dec 1963; LPW-3. FMNH 75617; (2, 59–71); Gulf of Mannar, Musal tivu; 7 Jan 1964; LPW-5.
Cocos-Keeling Islands: ANSP 128422; (13, 16–61); Direction I; 29 March 1974, Sta. 65.Thailand: ROM
70830; 2(67.2–73.0); Phuket, 7°74'02"N 98°24'45"E; 14 Mar 1997. ROM 71512; 15(10–27); Phuket,
7°36'31"N 98°22'28"E; RW 93–14; 16 Nov 1993. Indonesia: USNM 212858; (3, 47–59); Sumatra,
00°01'45''S, 98°31' 15''E; 0–1 m.; 25 November 1963. Savu Sea: UF 18358; (14, 29–66); Flores I. Australia:
Timor Sea: CAS 84078; (5, 31–38); Ashmore I.; 3 Jan. 1973. Pacific Ocean: South China Sea: Thailand:
Gulf of Thailand; CAS 84106; GVF 2185; (1, 81); Chumpon Prov.; 17 May 1960; Sta. 168. CAS 84020; GVF
2169; (1, 52) Koh Samet I.; 22 April 1960; Sta. 152. CAS 84039; GVF 1572; (2, 24–37); Koh Samet I.; 19
December 1957; Sta. 114. CAS 84003; GVF 2039; (1, 52); Koh Kra; 26 January 1960; Sta. 28. CAS 84540;
GVF 2172; (1, 52); Khorn Aho Bay; 24 Apr 1960. Vietnam: CAS 84097; GVF 2789; (5, 42–58); Hon Mieu
I.; 31 January 1961; Sta. 514. CAS 84054; GVF 2074; (3, 52–72); Nha Trang; 24 February 1960; Sta. 58.
CAS 84091; GVF 2796; (39 ,43–69); 2.8 mi. SSW of Nha Trang; 4 February 1961; Sta. 60-521. CAS 84087;
GVF 2116; (7, 48–78); Bay of Nha Trang; 14 March 1960; Sta. 60–99. CAS 95980; GVF 2113; (4, 54–60);
Ilot du Sud; 10 Mar 1960. China: Hong Kong: CAS 84046; GVF 1748; (76, 21–75); Pratas Reef; 23 May
1958; Sta. HK 71. CAS 95977; GVF 1745; (1, 58); Pratas Reef; 23 May 1958; Sta. HK 68. Taiwan: USNM
212845; (2, 13–57); Ta Fa Lieh; 27 May 1975. USNM 212846; (7, 42–74); Tan-Oz-Wan; 26 Dec 1969.
USNM 212847; (3, 42–43); Mao Bi Tou; 17 Dec 1969. Indonesia: FMNH 22036; (1, 25); Waigeo I; June
1929. SU 26650; (1, 44); Waigeo I.; 6 June 1929. Java Sea: CAS 35691; (1, 51); Java, N. of Jakarta, Pulau
Edam; 20 Dec. 1975. Molucca Sea; SMF 954–955 (2, 57–60) Ternate; 1894. MNHN 8697 (1, 56); Ternate;
Bleeker Coll. USNM 112062 (A24079); (1, 36); Moluccas, Tomahu I.; Albatross; 11 Dec 1908. SU 30017; (8,
40–67); Sangir Is., Sangi I.; 24 June 1929; A.W. Herre. Ceram Sea: USNM 261141; (9, 11–24);off Misool I.
Philippines: USNM 112057 (A14428); (1, 69); Luzon I., Cubat; Albatross; 23 Jun 1909. USNM 112056
Zootaxa 1691 © 2008 Magnolia Press · 17
status of Nectamia & 4 new species
(A23813); (1, 68); Luzon I., Biri Channel; Albatross; 2 Jun 1909. USNM 112197; (1, 60); Romblon Harbor;
25 Mar 1908. USNM 298560; (5, 31–63); Bantanes, 21°21'05" N. 121°55'14"E.; 31 May 1985. USNM
298657; (1, 23); Bantanes, Batan I.; GDJ 87-14; 2 May 1987. USNM 261153; (7, 14–58); Siquijor I.; SP 78-5;
10 May 1978. USNM 112060 (A23236); (1, 53); Tataan, Simaluc I.; Albatross; 20 Feb 1908. USNM 112053
(A18397); (1, 73); Cebu Mkt.; Albatross; 4 Apr 1908.USNM 112054 (A 23219-21); (3, 63–71); Mindinao I.,
Opal; Albatross; 4 Aug 1909. USNM 112052 (A6041-2); (2, 62–72); Tuminado I.; Albatross; 26 Feb 1908.
Sulawesi Sea: SU 27355; (6, 34–77); Jolo, Sulu I.; 1 Aug 1931. SU 33480; (9, 50–64); Sulu I.; 7 Jan. 1937.
SU 27356; (4, 19–73); Sulu I; 9 Aug 1931. USNM 112051 (A5310-12, 23701); (4, 62–75); Jolo I.; Albatross;
6–7 Mar 1908. Samar Sea: USNM 341630; (2, 65–74); Pangasinan; 14 Feb. 1980. USNM 341631; (4, 62–
72); Pangasinan; 14 Feb. 1980. Sulu Sea: ANSP 174727; (2, 56–57); Sibutu Island; 27 Apr 1926. CAS
98020; (37, 27–65); Layia, Luzon I.; 28 Jun 1948. CAS 53326; (4, 19–82); Dumaguete, Negros I.,; 10 Jul
1948. SU 33474; (27, 24–61); Dumaguete. CAS 84034; GVF 1614; (1, 63); Negros I; 20 Dec 1936. SU
27354; (16, 16–32); Negros I; June 1931; Herre Philippine Exp. A.W. Herre; USNM 261134; (27, 11–22); S.
tip of Negros I. USNM 261138; (55, 9–54); S. end Negros I. USNM 112050; (2, 35–65); N. Balabac Strait; 28
July 1958; Sta. 74. SU 27357; (14, 25–37); Zamboanga, Mindanao; 29 July 1931; A.W. Herre. USNM 112055
( A23637); (1, 65); Mindanao I., Murcielagos B.; Albatross; 21 Aug 1909. Philippine Sea: USNM 112058
(A23272); (1, 48); Samar I., Ft Palapag; Albatross; 2 Jun 1909. SU 40603; (1, 64); Samar I.; 10 Apr 1945; R.
Annereaux coll. Papua New Guinea: USNM 341632; (1, 44); Bougainville I.; TeVega Cr. 6; 19 Oct. 1984.
Coral Sea: USNM 261143, (11, 39–80). Lolorna I, Port Moresby. USNM 341625; (6, 32–68); Louisade
Arch., Panapompom I.; BBC-1693, 15 June 1979. Australia: Coral Sea: ANSP 123319; (2, 27–27); Big
Hope Island; TSA-34; 27 Jan 1969. Endeavour Reef: ANSP 123377; (3, 19–24); TSA-13; 13 Jan 1969. ANSP
123348; (1, 19);TSA-12; 13 Jan 1969. ANSP 123440; (5, 25–61); TSA-17; 16 Jan 1969. CAS 28511; (2, 41–
42); Wharton Reef; 9 October 1960; SI0-61-132; CAS 28519; (1, 60); Flinders I; May 1966; SI0 69-222;
USNM 212868; (10, 47–74). One Tree I.: ANSP 135401; (7, 31–45); DFH 72-79; 5 Oct 1972. BPBM 14421;
(2, 34–64); USNM 212867, (1, 32); USNM 212871, (2, 46–75); USNM 212869; (38, 36–73); USNM 212870;
(34, 43–77); USNM 213125; (1, 60); BBC 1407. USNM 177036; (10, 36–52). Green I, near Cairns. New
Caledonia: Coral Sea: BPBM 33703; (4, 38–72); Îles Chesterfield, Ilot du Mouillage; color largest; 28 Aug.
1988. BPBM 33532; (2, 54–60); Color smaller; Îles Chesterfield; 6–9m.;20 Aug. 1988. Solomon Sea: Trobri-
and I.: USNM 212843; (1, 28). USNM 213232; (1, 41); 11 Jun 1970. Bismark Archipelago: USNM 261142;
(9, 16–23); Umboi I., 6°41'54"S, 147°53'48"E. Solomon Islands: FMNH 22031; (1, 63); Ugi I; 10 Apr. 1929;
Crane Pacific Exp. BPBM 15628; (1, 45); Florida I.; 21 July 1973. Santa Cruz Islands: CAS 84103; GVF
1817; (1, 44); Vanikoro I.; 1 Oct. 1958; Sta. HK138. USNM 357434; (2, 44–66); 10°17'S 166°18'30"E; 15
Sep 1998; 0–1.5 m. Tonga Islands: USMN 341610; (4, 42–47); JTW 93-16; 2Nov. 1993. Ryukyu Islands:
USNM 72562; (8, 18–42); Luchu I., Naha, Okinawa; Albatross. USNM 132681; (1, 39). USNM 212854; (1,
46); 22 Nov 1964. USNM 212855; (8, 41–70); Caroline Islands: Palau: Angaur I.: CAS 84089; GVF 1425;
(60, 25–73); 22 Oct 1957; Sta. 45. CAS 84014; GVF 1423; (2, 27–46); 21 Oct. 1957; Sta. 43. MCZ 29742; (4,
24–33);1862. CAS 84105; GVF 947; (1, 33); Kayangel I; 8 Oct 1956; Sta. 161. CAS 4019; GVF 762; (3, 35–
43); Eil Malk Lagoon; 27 Oct. 1955; Sta. 252. CAS 84096; GVF 902; (2, 28–30); Peleliu I.; 19 Sept. 1956;
Sta. 116. CAS 84102; GVF 1852; (1, 53); Aurapushekaru I.; 20 Jan 1959; Sta. 24. CAS 84027; GVF 798; (3,
44–51); Auluptagel I.; 8 July 1956; Sta. 12. CAS 84017; GVF 1399; (1, 22); Chol I; 23 Sept. 1957; Sta. 21.
CAS 95979; GVF 1829; (1, 29); Babelthaup I.; 23 Jun 1958; Sta 58-1. CAS 84101; GVF 1865; (1, 53);
Ngercheu I.; 4 Mar 1959. CAS uncat.; (4, 19–49); 12–18 Oct 1933. Kapingamarangi Atoll: CAS 84021; (6
spec); 1°3'27"N; 154°46'52"E; GVF 338; 12 Jul 1954. CAS 84032; GVF 462; (58, 10–30); 16 Aug 1954; Sta.
159. CAS 84008; GVF 317; (12, 29–49); 1 July 1954; Sta. 14. CAS 84030; GVF 352; (2, 33–34). CAS 84036;
GVF 354; (13, 10–57). CAS 84037; GVF 386; (25, 10–48). CAS 84040; GVF 367; (10, 32–56). CAS 95970;
GVF 475; (3, 27–31). CAS 95973; GVF 446; (6, 24–39); Ya p I.: CAS 84099; GVF 1918; (11, 29–63); 23 Dec
1959; Sta. 95. CAS 84011; GVF 1923; (1, 51); 28 Dec 1959; Sta. 100. CAS 84042; GVF 1907; (4, 42–69); 13
Dec 1959; Sta. 84. CAS 84093; GVF 1919; (21, 39–70); 24 Dec 1959; Sta. 96. Pohnpei: CAS 84098; GVF
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18 · Zootaxa 1691 © 2008 Magnolia Press
496; (6, 16–50); Na I.; 6 Oct. 1954; Sta. 193. CAS 95978; GVF 492; (15, 16–55);1 Oct. 1954; Sta. 189. CAS
95971; GVF 499; (7, 17–26); 8 Oct. 1954; Sta. 196. Ifalik Atoll: CAS 84104; GVF 122; (8, 31–54); 13 Sept.
1953; Sta. 1. CAS 84009; GVF 133; (1, 40); 23 Sept. 1953; Sta. 12. CAS 84100; GVF 249; (16, 22–38); 26
Oct. 1953; Sta. 128. CAS 83796; GVF 262; (3, 52–59); 28 Oct. 1953; Sta. 141. CAS 95972; GVF 185; (7, 26–
63); 8 Oct. 1953; Sta. 64. SU 29513; (2, 33–47). Rug I.; A.P. Landon coll. Kosrae: MCZ 29503; (1, 23).Mar-
shall Islands: Arno Atoll: USNM 166597; (3, 21–38). Bikar Atoll: USNM 212848; (3, 39–43). Bikini Atoll:
ANSP 91322; (1, 48); Namu I.; 3 Apr 1946. FMNH 60021; (9, 43–70); Bikini I.; 22 Mar. 1946. USNM
112352; (10, 19–72); Romuk I.; x-ray. USNM 112351; (20, 45–65); 1 Apr 1946. USNM 112353; (5, 49–68);
17 Apr 1946. USNM 166597; (2, 32–36); 22 Jun 1950. Enewetak Atoll: BPBM 8860; (1, 50); 2 Jan. 1970.
FMNH 60022; (3, 58–67); Aaraanbiru I; 3 June 1946. UF 42278 (12, 16–71). USNM 112347; (121, 15–
72);Jul 1950. USNM 112348; 52 spec; S-56-197; 1 Jul 1946; (1, 62.3) cleared & stained. Kwajalein Atoll
USNM 212851; (1, 40); 8 Nov 1964. Likep Atoll: USNM 152947; (1, 59); 20 Aug 1949. Rongelap Atoll:
FMNH 60023; (9, 17–39); Naen I.; 30 July 1946. USNM 112350; (17, 23–66); 19 Jun 1946; x-ray. Rongerik
Atoll: USNM 112349; (3, 28–61); 14 Aug 1947. Mariana Islands: Guam: ANSP 174728; (3, 36–48); AS00-
G-6-1; 1 Jun 1900. BPBM 97; (1, 70). 14 Jun 1900. SU 19280; (2, 29–36); 15 Aug 1952. CAS 84007; (1, 44),
CAS 95974; GVF 1839; (9, 26–44); 10 Oct 1958; Sta. 11. CAS 84088; GVF 1842; (12, 25–57);19 Oct. 1958;
Sta. 14. CAS 84018; GVF 1840; (26, 71); 12 Oct. 1958; Sta. 12. CAS 84090; GVF 1859; (96, 15–55); 7 Apr.
1959; Sta. 31. CAS 84086; GVF 1874; (45, 10–60); 26 Apr. 1959; Sta. 46. CAS 84035; GVF 1845; (1, 32); 9
Jan. 1959; Sta. 17. CAS 84026; GVF 1045; (2, 28–34); 1956; Sta. 248. USNM 139811; (41, 24–66); 26 Nov
1945. USNM 154243; (31, 18–66). UF 117138; (1, 47); 1995. USNM 124024; (7, 17–27); 7Jul 1945. USNM
124135; (1, 55); 20 May 1945. USNM 212856; (13, 56–89); N. end of Tumon Bay. USNM 139799; (3, 17–
22); 12 Nov 1945. UGM 4183; (1, 43); North of NCS beach, HK0105; 5 Nov. 1967; UGM 4716; (12, 18–58);
Tumon Bay, Reef Flat; 14 July 1970; UGM 1420; (1, 66.4); Togcha Bay, GFW-117; 27 Oct. 1964; UGM
1433; (1, 68.1) Togcha Bay, GFW-116; 27 Oct. 1964. Saipan: ANSP 119642; (6, 23–41); SAI 1-47; 24 Jul
1954. ANSP 119418; (1, 40); SAI 17-49; 27Aug 1954. ANSP 152653; (1, 65); SAI 1-25; 18 Oct 1953. ANSP
152626; (8, 39–79); SAI 1-48; Aug 1954. ANSP 152649; (4, 39–48); SAI 1-48; Aug 1954. ANSP 152622;
(12, 41–77); SAI 1-48; Aug 1954. ANSP 152632; (9, 44–73); SAI 1-48; Aug 1954. ANSP 152657; (1, 74);
SAI 17-49; 27 Aug 1954. SU 29156; (4, 27–53); 7 Dec. 1933; A.W. Herre. CAS 84103 GVF 789; (1, 55); 16
June 1956; Sta. 3. USNM 123828; (3, 20–31) Jun 1945. USNM 150008; (20, 20–33); 19 Dec 1948. USNM
154242; (2, 43–51); Jul 1949. Tinian: SU 29158; (74, 20–67); Tinian I.; 17 June 1956; Sta. 4. FMNH 47315;
(10, 32–48); 7 Oct 1933. Samoan Islands: American Samoa; Tutuila: Pago Pago: ANSP 51490; (14, 30–
49); CFS17-PP-S; 1917. ANSP 51504; (5, 32–49); 17-4-5-PP; 5 Apr 1917. CAS 82687; (4, 61–66); Pago
Pago; 14 Nov 1946. SU 67429; (18, 20–56); 1902. USNM 112048; (11, 22–60); U.S. Fish Tag 04608. USNM
114973; (3, 40–63); 3 Jun 1939. USNM 212836; (3, 21–55); 19 Aug 1963. USNM 212837; (1, 35). Western
Samoa: Upolu: Apia: ANSP 151657; (8, 24–47); DSJ-APIA. CAS 95981; (out of IUM 10894); (5, 23–51);
1902. MCZ 50056; (7, 22–61); 1902; U.S. Fish tag 06556. USNM 112047; (11, 26–58); 1902. USNM 11966;
(3, 23–46); 1902. USNM 212838; (4, 27–44); 1902. Fiji: ANSP 89299; (1, 24); Che33-F-2; Feb 1933. CAS
220972; (1, 65); Budd Reef, Mbeka I., 16°30.444'S, 179°40.571'W; G03-54; 22 May 2003; 1.2 m. Nukulau I.:
CAS 218295; (4, 13–26); 18°10.726'S, 178°30.799'E; G02-17; 28 Jan 2002; 0.1 m. FMNH 22007-028; (22,
25–54); 15 Mar 1929; A.W. Herre; Crane Pacific Exp.; SU 24934; (19, 20–41); 16 Mar 1929; A.W. Herre
(Crane Pacific Exp). Viti Levu : CAS 7452; (1, 57); Suva; Mar. 1932; W. Cassin coll. CAS 218010; (1, 38);
off Suva, 18°10.399'S, 178°25.277'E; G02-50; 10 Feb 2002.CAS 218012; (16, 19–43); off Suva, 18°11.227'S,
178°31.040'E; G02-04; 23 Jan 2002; 1.2–1.5 m. CAS 218014; 5, (15–33); off Suva, 18°11.227'S,
178°31.040'E; G02-03; 23 Jan 2002; 1 m. Vanau Balavu: CAS 223741; (6, 24–61); Balavu Harbor,
17°11.212'S, 179°00.095'W; G02-03; 8 Jan 2003. Rotuma: USNM 288051; (7, 32–58); VGS 86-3; 8 May
1986. USNM 212872; (2, 27–27); 12°30'00"S; 177°00'00"E. Vanuatu: ANSP 85882; (1, 55); Efate. ANSP
91561; (1, 31); Anelganket; ML53-AA; 1953. FMNH 22030; (1, 53); Malo I.; 30 Mar. 1929. SU 25089; (2,
49–56), Malo I.; 30 Mar. 1929. FMNH 22029; (1, 55); Vila, Efate; 27 Mar. 1929; Crane Pacific Exp. SU
Zootaxa 1691 © 2008 Magnolia Press · 19
status of Nectamia & 4 new species
25001; (2, 41, 53); Vila, Efate; 27 Mar. 1929; A.W. Herre coll. BPBM 5782; (2, 52–58); Efate; 25 Nov. 1964.
Kiribati: Phoenix Islands: USNM 167316; (5, 37–82); Onotoa I; 2 Dec. 1954. USNM 114972; (76, 36–62);
USNM 114975; 14(42–64); Kanton I. Lagoon. USNM 114974; (47, 8–42); Hull I.; 8–12 Jul 1932. USNM
114975; (14, 43–64); Kanton I. Lagoon; 25–26 May 1939. USNM 179689; (1, 29); Kanton I.; 26 May 1939.
Tokelau Islands: USNM 212840; (28, 12–28); Atafu I.; 27 May 1975. USNM 212839; (2, 56–57); Fakaofo
Atoll.
Diagnosis. A Nectamia with a few faint or no pale bars on body, basicaudal spot diffuse in adults, caudal
fin without dark margins, subocular mark narrow, linear (Tab. 2); usually 13 pectoral-fin rays; total gill rakers
usually 24–28, 25–28 in the Indian Ocean and 24–27 in the Pacific Ocean (Tab. 5); body depth 37–42%, cau-
dal-peduncle depth 15–18%, second anal-spine length 15–17%, and pectoral-fin length 24–26% of standard
length (Tab. 3).
Description. For general body shape see Table 3 for range of proportions (as percentage of standard
length) with the neotype first and additional material in parentheses.
Dorsal fin VII(I)–I,9 (Fig.1A); anal fin II,8; pectoral-fin rays usually 13-13, rarely 14-13; pelvic fin I,5;
principal caudal rays 9+8; pored lateral-line scales 24; transverse scale rows above lateral line 2; transverse
scale rows below lateral line 5(5–6); median predorsal scales 3–4; circumpeduncular scale rows 12 as 5+2+5;
total gill rakers 27 (23–29), well developed 25(20–25), upper arch 2+4 (1-3+4-6), lower arch 20+0 (17-20+0-
1), second arch with shorter robust gill rakers 4+18 (Fig2. C and D).
Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and
vomer; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10+14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural.
Three supraneurals, two supernumerary spines on first dorsal pterygiophore. Basisphenoid present. Supra-
maxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal mar-
gins. Infraorbital shelf present on third bone. Scales ctenoid on cheek, subopercle, opercle, nape, isthmus,
cycloid on base of pectoral fin and ctenoid behind pectoral fin and rest of body. Pored lateral-line scales com-
plete, simple with single pores extending above and below main canal (Fig. 4A). No axillary scale at base of
pelvic spine (Fig 3A). Two large ctenoid scales on base of pelvic fins.
Cephalic lateralis pores mostly minipores (Fig. 5), a terminal lachrymal pore, a pair of ventral lachrymal
pores, an anterior dentary and mental pores, terminal end of supraorbital canal with a large pore near anterior
nare, lateral margin of supraorbital canal near posterior nare small, lateral margin of supraorbital canal mid-
way on interorbit with small pores, simple postorbital canal projections with minipores, anterior margin of
supratemporal canal smooth, posterior supratemporal canal margin with complex projections and minipores,
lateral margin of mandibular canal with many minipores, a small posterior articular pore.
Life colors. See Figure 6H. Kuiter and Kozawa (1999) provide a number of underwater photographs (p.
32, A–E) identified as Apogon guamensis. Their Apogon species 16 (p33, A–B) may be this species. Fowler
and Bean (1930, p. 44) gave field color notes recorded by others for Albatross specimens USNM 112060
(A23236) and USNM 112168 (A232337) however two species were involved (Nectamia fusca and N. savay-
ensis). Head, body and fins without stripes; head with dark narrow cheek mark to near angle of preopercle,
lachrymal and anterior infraorbital with yellowish mark, iris yellowish with white inner ring; head and body
bronze to brownish background sometimes with faint lighter bars; caudal peduncle with a diffuse basicaudal
spot mostly above pored lateral-line scales, more intense and discrete in young and juveniles; first dorsal fin
with dusky membranes between spines II–IV, second dorsal, pectoral, pelvic anal and caudal fins pale.
Preserved color pattern. Adults (Tab. 2): narrow, linear subocular cheek mark behind upper jaw; first
dorsal fin with dark membrane between spines 1–4; edges of caudal lobes plain; caudal peduncle with a dif-
fuse spot of variable intensity; pale bars, usually four or fewer, occasionally present on body from below the
first dorsal fin and behind the pectoral fin to just behind the ends of the second dorsal fin base and anal fin
base, bars extend above the lateral-line scales starting about the second dorsal fin; no dark saddle under the
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20 · Zootaxa 1691 © 2008 Magnolia Press
first or second dorsal fin; stomach and intestine black; roof of mouth pale to scattered tiny melanophores not
extending onto gill arches. Juveniles: similar to the adults except basicaudal mark more intense and spot-like.
FIGURE 9. Distribution of material examined for Nectamia fusca .
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status of Nectamia & 4 new species
TABLE 5. Geographic variation in gill-raker and rudiments counts for Nectamia fusca.
* = neotype
N=249 Upper Arch Lower Arch Total
5678x 17 18 19 20 21 x 23 24 25 26 27 28 29 x
RED SEA
Israel 1 1 1
Egypt 737.3 171119.2 1522 26.5
Saudi Arabia 31226.9 376119.3 21841126.2
Ethiopia 1 1 1 6.0 1 2 18.7 2 1 24.7
INDIAN OCEAN
Kenya127.7 319.0 1226.7
Tanzania/Zanzibar 1517.0 241 18.9 1132 25.9
Mozambique 317.3 22 18.5 211 25.8
Madagascar 1 1 6.5 1 1 18.5 1 1 25.0
Maldives 12 14 6.5 2 5 18 1 19.7 4 13 8 1 26.2
Cocos-Keeling Is 2 3 6.6 3 2 19.4 1 3 1 26.0
Ashmore I. 32 6.4 1211 18.4 1112 24.8
Indonesia 46 6.6 163 18.2 1333 24.8
PACIFIC OCEAN
Ryukyu Is. 5 2 6.3 3 4 18.6 3 2 2 24.9
Hong Kong 8 5 6.4 5 8 18.6 4 5 4 25.0
Vietnam 21316.9 375119.3 13561 26.2
Thailand 2216.8 5 19.0 221 25.8
Philippine 610 6.6 196 18.3 1465 24.9
Indonesia 3 7.0 2 1 18.3 2 1 25.3
Guam 7* 27 1 6.8 3 26 6* 19.1 1 8 20 5* 1 25.9
Eniwetok Atoll 2816.9 1262 18.8 11153 25.7
Rongelap Atoll 3 1 6.3 4 19.0 3 1 25.3
Bikini Atoll 2 4 6.7 2 3 1 18.8 4 1 1 25.5
Australia 5 6.0 14 18.8 14 24.8
Fiji 4416.7 153 19.2 4311 25.9
Samoan Is141 6.0 15 18.8 231 24.8
Vanuatu 31 6.3 13 18.8 121 25.0
Canton I. 97 6.4 3121 18.9 2851 25.3
SUMMARY
Red Sea / East Africa 1 6 30 10 7.0 10 27 8 2 19.0 6 5 20 12 3 1 26.1
East Indian Ocean 21 25 6.5 2 10 12 21 1 19.2 2 4 9 21 9 1 25.7
China Sea 17 22 2 6.6 11 24 5 1 18.9 8 12 13 7 1 25.5
Philippines / Indonesia 6 13 6.7 1 11 7 18.3 1486 25.0
Guam / Marshall Is 14 40 2 6.8 1 7 39 9 19.0 1 2 16 27 9 1 25.8
Coral Sea / Samoa1251316.4 7294 18.9 621102 1 25.3
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22 · Zootaxa 1691 © 2008 Magnolia Press
Distribution. Known from the Red Sea, Indian Ocean (except the Seychelles, Chagos Archipelago, Car-
gados Carajos, Mauritius, Reunion and Rodrigues) and throughout the West Pacific to the Tonga Islands,
Samoa, Phoenix and Marshall Islands (Fig. 9).
Status of Apogon fuscus. Quoy and Gaimard described Apogon fuscus collected at Guam in 1824 during
the expedition of L. de Freycinet from the 1817–1820 voyage around the world by the Uranie and Physici-
enne.
No type material was listed by Bauchot and Desoutter (1987) for Apogon fuscus at the Muséum National
d'Histoire Naturelle. Their discussion about additional type material of Apogon guamensis for the material in
MNHN 8682, collected at Guam during the 1817–1820 expedition was inconclusive. One specimen in
MNHN 8682 is about 69 mm total length, the other two are 70 and 75mm total length. All specimens are iden-
tified here as conspecific with Apogon guamensis in agreement with Bauchot and Desoutter. A single speci-
men, about 2.5 inches (~68 mm) total length, was mentioned by Quoy and Gaimard in their description of
Apogon fuscus. The 52.5 mm SL specimen in MNHN 8682 is almost identical in total length. This specimen is
treated as the neotype for Apogon fuscus. It may have been the basis of the original description, but its status
as the holotype cannot be determined. Curiously, this earlier description and material was not discussed by
Valenciennes when describing Apogon guamensis in 1832 from the expedition of J. Dumont d’ Urville of
1826–1829, also from Guam.
The description generally fits any of three species known from collected material to inhabit the waters of
Guam. These species are: Nectamia fusca, N. savayensis, N. bandanensis The species most commonly col-
lected over time from Guam has been N. fusca. The other two species are relatively rare in collections from
Guam, but may have been more abundant in the past. One other new species, Nectamia luxuria should occur
at Guam because it has been found at Yap and Saipan.
Historical use of Apogon fuscus has been inconsistent. Jordan and Seale (1906) erroneous identified and
illustrated a six-spined species as Amia fusca. Later, when Jordan (1917) erected Nectamia as a genus based
on Apogon fuscus, he briefly described the species as “...differs from the type of the genus in the long and
rounded caudal fin, attached to a long caudal peduncle as long as head. It has large scales and six dorsal
spines.”. Kuiter and Kozawa (1999, p. 43) identify a similar six-spined species and appear to have followed
Jordan and Seale. Quoy and Gaimard (1824) clearly described a seven-spined species “La première dorsale a
sept aiguillons; la seconde, un seul, et dix rayons branchus...”. Fowler and Bean (1930) treated the species a
valid seven-spined species, but in another group with fewer gill rakers, stripes along the base of the dorsal fins
from the nape and just above the eye to upper surface of the caudal peduncle and a small basicaudal spot.
Apogon fuscus was not used or discussed by Lachner (1951). Apogon fuscus has been used more frequently
since 1990 (see Eschmeyer et al. 2007). Greenfield (2001) concluded that Jordan and Seales’ use of the name
was a misidentification as was Kuiter and Kozawa’s use. The illustration by Jordan and Seale (1906, p. 244,
fig.38) is a new species of Apogon, A. seminigracaudus Greenfield (2007).
Nectamia is based on Apogon fuscus, and without fixing the generic name, its use as a genus or subgenus
would have an uncertain status. Gon (1987) fixed a species name for the poorly described genus Ostorhinchus
Lacépède, 1801, as a cardinalfish, Ostorhinchus fleurieu. This genus was the senior synonym for the species
including Nectamia. Apogon aureus, a close relative of A. fleurieu (Randall et al. 1990), and N. fusca, as
Apogon guamensis, were not associated in the same broad clade recognized by Mabuchi et al. (2006, figs. 2
and 3). Apogon fucus is treated as the senior synonym of Apogon guamensis, an unavoidable disruption of
existing usage. Nectamia is treated as a valid genus.
Remarks. This species has an unusual distribution in the Indian Ocean, not reaching islands on the Mas-
carene Plateau. Slight geographic differences in the gill-raker counts occur for the major oceans and data
reported here extends Lachner’s (1951) observation about Red Sea material to the western Indian Ocean for a
slightly higher number of gill rakers. The range and frequency of gill rakers appear to be slightly lower in the
Indonesian-Philippine region. No clear differences were observed in the preserved color patterns or from
Zootaxa 1691 © 2008 Magnolia Press · 23
status of Nectamia & 4 new species
color underwater slides to suggest that two species may exist. Nectamia fusca is a slender species with a
uniquely narrow, oblique cheek mark below the eye, slightly shorter pectoral-fin length, well-developed cau-
dal spot in juvenile becoming diffuse in adults unlike the other eight species.
This species has been collected at stations with one or more of the following species: Nectamia savayensis
(ANSP 85880, 98137) and N. luxuria (CAS 28505, 52558, FMNH 22032, USNM 52432). A number of spec-
imens were found with buccal enlargement: eggs present for USNM 212868, 1 specimen, 68 mm SL, no eggs
were present (USNM 112046, 112197, 147524, 212845, 212858, 212846, 212868, 167316 and CAS 53326,
84020, 84054, 84087, 84091). Apogon ocellatus and Apogon spongicolus are juvenile specimens of Nectamia
fusca. The largest of 2285 specimens was 80 mm SL.
Nectamia ignitops new species
Figures 3F, 8, 10, 18 and Tables 2–3, 6
Type material: Holotype BPBM 21897 68.1 mm SL; Malaysia, Tioman I.; J. E. Randall; 2 Aug. 1977; color
photograph; x-ray. Paratypes BPBM 38585; (4, 61.1–75.0); same data as holotype; x-ray.
Additional material: Thailand: CAS 84064; GVF 2168; (2, 50–75); Parsho Bay, Rayong Prov.; 21 Apr
1960. CAS 82649; GVF 2646; (7, 53–73); Prachuap Khiri Khan; 17 Jun 1961. CAS 82650; GVF 2183; (5,
54–79); off Koh Kroi I.;25 Jun 1968; 15–60 m. Vietnam: Nha Trang Bay: ROM 73558; (2, 63–63); Hon Mot,
12.1755°N, 109.2781°E; RW02-01; 11 May 2002; 7 m; color photos. ROM 79173; (3, 58–71); Hon Tom,
12.744°N, 109.23944°E; RW02-20; 13 Jun 2003; 2–4 m. ROM 79174; (27, 57–73); Hon Lon, 12.1822°N,
109.29361°E; RW02-05; 13 Jun 2003; 4–8 m. ROM 79175; (20, 15–79); Hon Mot, 12.17694°N,
109.27889°E; RW02-19/21, mixed; 13 Jun 2003; 0–6 m.
Diagnosis. A Nectamia with two faint dorsal saddles, faint caudal-peduncle band, no pale bars on side of
body, and faint cheek mark, caudal fin without dark margins (Tab. 2, Fig. 8); 13 pectoral-fin rays; total gill
rakers usually 24–27 (Tab. 6); body depth 44–47%, caudal-peduncle depth 18–20%, second anal-spine length
19–21%, and pectoral-fin length 25–29% of standard length (Tab 3).
Description. For general body shape see Figure 10. Range of proportions (as percentage of standard
length) in Table 3 with paratypes and non-type material in parentheses.
Dorsal fin VII(I)–I,9; anal fin II,8; pectoral-fin 13-13; pelvic fin I,5; principal caudal rays 9+8; pored lat-
eral-line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line 6;
median predorsal scales 2; circumpeduncular scale rows 12 (5+2+5); total gill rakers 25 (24–28), well devel-
oped 24 (21–25), upper arch 1+6 (0-2+5-6), lower arch 18+0 (17-19+0-1).
Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and
vomer; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10+14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural.
Three supraneurals, two supernumerary spines on first dorsal pterygiophore. Basisphenoid present. Supra-
maxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal mar-
gins. Infraorbital shelf present on third bone. Ctenoid scales on cheek, subopercle, opercle, nape, isthmus,
cycloid at base of pectoral fin, ctenoid on rest of body, base of pelvic with two large ctenoid scales, no axillary
scale present. Pored lateral-line scales complete, fourth scale variably complex, holotype with two large pores
and three small pores above main canal and one large and two small pores below (Fig 3F).
Etymology. A combination from the Latin ignitus meaning glowing (of a fire) and the Greek ops meaning
eye, referring to the color of the iris. Treated as a noun in apposition.
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24 · Zootaxa 1691 © 2008 Magnolia Press
FIGURE 10. Nectamia ignitops, holotype, BPBM 21897, 68.1 mm SL, Malaysia, Tioman I., J. E. Randall, 2 Aug. 1977,
6 m.
TABLE 6. Geographic variation in gill-raker and rudiments counts for Nectamia ignitops.
* = holotype
Life colors. Unknown from underwater photography. From photograph of freshly collected: holotype
with a reddish iris; silvery on the side of head and body, a faint subocular cheek mark tannish dorsally, without
dark saddles, pale band on anterior caudal peduncle; blackish membrane (distal two thirds) between the third
and fourth first dorsal spines, a lighter dusky membrane between the fourth and fifth spines, membranes
dusky (darker distal two thirds) between the fifth and seventh spines and dusky between the second and third
anal spines; second dorsal fin, anal fin and caudal fins pale.
Preserved color pattern. Holotype (Fig. 10, Tab. 2): faint subocular cheek mark as a faint line of melan-
ophores; first dorsal fin without dark membrane between spines 1–4; edges of caudal lobes plain; caudal
peduncle without obvious dark bar or spot; no obvious dorsal saddles under the first or second dorsal fin, paler
bar-like area on anterior caudal peduncle; stomach and intestine black; roof of mouth and gill arches pale.
Paratypes: similar to holotype except smallest paratype with traces of the dorsal saddles and darker band.
Juveniles: unknown. Vietnam specimens with a narrow cheek mark, first dorsal fin with dark membrane
between spines 1–4; edges of caudal lobes plain; dorsal dark saddles under the first or second dorsal fin; cau-
dal peduncle usually with a complete dark bar; pale area between peduncle bar and second dorsal saddle;
stomach and intestine black; roof of mouth blackish with melanophores extending down onto lower gill arch
Distribution. Known from coastal South China Sea (Fig. 8).
Remarks. Recent material (BPBM 21897) from Tioman Island, Malaysia was initially identified as
Apogon bandanensis, however, no cheek mark, saddles, or caudal mark were present in the fresh photographs
and faint or absent on the preserved holotype and paratypes. A communication from J. E. Randall suggests
that the color pattern may have faded prior to photography. Icing fish prior to photography may affect melan-
N=41 Upper Arch Lower Arch Total
678x 17 18 19 20 x 24 25 26 27 28 x
Tioman I. 5* 7.0 3* 1 1 18.6 3* 1 1 25.6
Gulf of Thailand 2 10 2 7.0 4 8 2 17.9 5 6 3 24.9
Vietnam 11 11 7.5 1 4 14 3 19.0 1 2 9 8 2 26.5
Total 2 26 13 7.3 5 15 17 4 18.5 6 11 13 9 2 25.6
Zootaxa 1691 © 2008 Magnolia Press · 25
status of Nectamia & 4 new species
ophore pigment patterns. These specimens were taken with Nectamia luxuria (BPBM 21896). Four other lots,
CAS 82649, 82650, 84058, 84064 from Thailand are very similar to the Tioman specimens and are tentatively
identified as this species. Several of these Thailand specimens had faint cheek marks and two specimens had a
faint caudal-peduncle mark. Additional material from Vietnam (ROM 79173, 79174, 79175) is tentatively
identified as this species, Some of this material was similar in color pattern to the Thailand specimens, but one
lot suffered from partial dessication and station mixing. None of the other Vietnam specimens examined had
bold contrasting color patterns in preservation. Most specimens had a complete caudal band. This species usu-
ally has 24–27 total gill rakers versus 26–28 total gill rakers in Nectamia bandanensis. The largest of 69 spec-
imens was 79 mm SL.
FIGURE 13. The distribution of collection sites for Nectamia zebrinus () N. savayensis with regional total gill raker
averages.
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26 · Zootaxa 1691 © 2008 Magnolia Press
Nectamia luxuria new species
Figures 3E, 6B, 11–12, 18 and Tables 2–3, 7
Type material: Holotype USNM 209644 59.1 mm SL; Indonesia, Ceram, Tandjung Liang; 10 Jan 1973;
Rumphius Exp. I, Sta. Li-1; VGS 73-6. Paratypes USMN 344924 (15, 23–61); Same data as the holotype.
USNM 213235; (7, 55–65); Pulo Siburu, Mentawei Is; 30 Nov 1963; Te Vega Sta. 104. USNM 213342; (24,
19–56);Borneo, Celebes Sea, Darvel Bay; 2 Feb 1965; Te Vega Cr. 6 Sta. 216. ROM 54318; (3, 45–61);
Negros Oriental, Siquijor I.; RW 87-8; 9 May 1987. USNM 213159; Australia, One Tree I., (7, 38–68); 25
Nov 1966; VGS 66-8. BPBM 21896; (9, 41–77); Malaysia, Tioman I.; 2 Aug. 1977.
Additional material: Indian Ocean: Cocos-Keeling Islands: ANSP 128423; (2, 35–57); Pulo Pandang;
17 Mar 1974; Sta. 40. Maldive Islands: CAS 95967; (2, 43–51); Male Atoll, Hulele I.; Te Vega 64-5; 8 Nov
1964. CAS 95984; (26, 33–53); Male Atoll, Hulele I.; 8 Nov 1964. Thailand: ROM 71511;13(14–71);
Phuket, 7°47'10"N 98°23'45"E; RW93-03; 0–1m. ROM 71528; 8;20–69; Phuket, 7°45'47"N 98°20'24"E; 10
Nov 1993; 0–6m. USNM 213151; (8, 23–63); Koh Huyong, Similan I; 3 Nov 1963; Te Vega Sta. 78. Indone-
sia: Off Western Sumatra: USNM 213153; (19, 19–58); Pulo Bai, Batu Group; 25 Nov 1963; Te Vega Cr H,
Sta. 101. USNM 213235; (7, 55–65); Pulo Siburu, Mentawei Is; 30 Nov 1963; Te Vega Sta. 104. Pacific
Ocean: China: Hong Kong: SU 61220; (14, 55–70); Pratas Reef; 26 May 1958; Sta. HK 77. GVF 1745; (2,
52–53); Pratas Reef; 23 May 1958; Sta. HK 68. CAS 98021; GVF 1750; (5, 59–68); Pratas Reef; 25 May
1958; Sta. HK 73. Thailand: CAS 84154; GVF 1535; (8, 45–52); Koh Tao I.; 10 Nov 1957. ROM 71511; (13,
14–71); Malacca Strait, Phuket, 7°47'10"N 98°23'45"E; RW93-03; 0–1m. ROM 71528; (8, 20–69); Malacca
Straits, 7°45'47"N 98°20'24"E; 10 Nov 1993; 0–6m. Vietnam: CAS 85997; GVF 2116; (3, 48–58); Bay of
Nha Trang; 14 March 1960; Sta. 60–99. Malaysia: Sabah: USNM 213342; (24, 19–56); Darvel Bay; 2 Feb
1965; Te Vega Cr. 6 Sta. 216. Indonesia: CAS 98022.; (3, 47–54); Sangi Siang I.; JEM 87-7; 24 Sep 1987.
Banda Sea: USNM 209644 (1, 59); Ceram, Tandjung Liang; 10 Jan 1973; Rumphius Exp. I, Sta. Li-1; VGS
73-6. USNM 210243; (5, 24–53) Ambon I.; 8 Jan. 1973; Rumphius Exp. 1, Sta. AB-3; VGS 73-20. BPBM
18537; (1, 34); color, black & white; Ambon I., Eri; 6 m; 31 Jan. 1975. USNM 341617; (8, 40–55); Nusa
Laut; 16 Jan 1973; Rumphius Exp. I, Sta. NU-1. USNM 210379; (2, 53–58) Saparua; 20 Jan 1973; Rumphius
Exp. I, Sta. Kw-1; VGS 73-16. USNM 341629; (1, 34); SE Naira I, Banda I.; 8 Mar 1974. VGS 74-9. Makas-
sar Strait: BPBM 26723; (2, 43–56); Sulawesi, Bonebetang I.; 1.5m; 8 Sep. 1978. Molucca Sea: USNM
171396; (A.23613); (1, 51); Jillolo, Halmahera I; 1 Dec. 1909. USNM 171398 (A23883); (1, 41); USNM
171394 (A.23529); (1, 73); Tidore I; 25 Nov 1909. USNM 112237 (A23527, 23528, 23530, 23884) (4, 63–
77); Tidore I, S of Ternate; 25 Nov 1909. USNM 112235 (A23484); (1, 58); Kayoa I; 29 Nov 1909. USNM
171383 (A.23105, 106); (2, 49–53); Tomahu I; 11 Dec 1909. USNM 171400 (A.24061); (1, 51); Talisei I, N.
of Sulawesi; 9 Nov 1909. USNM 112193 (A.14342); (1, 49); Tobea I.; 14 Dec. 1909. Irian Jaya: FMNH
22037; (1, 24); Waigeo; June 1929. Philippines: ANSP 77377; (8, 41–51); CMP-P; Nov 1917. ANSP 48885;
(8, 43–52); CMP-P; Nov 1917. USNM 341621; (7, 37–60); Sangay Siapo I.; EM 88-9; 28 May 1988. USNM
268261 (3, 57–62) and 341620; (1, 67); Negros Oriental, Siquijor I.; LK 79-15, 15 May 1979. USNM 341622;
(1, 41); Negros I., south tip; SP 78-9; 12 May 1978. ROM 54321; (9, 9–56); Cebu, Similon I.; RW87-31; 20
May 1987. ROM 73808; (8, 49–56); Cebu, Similon I., Rohol Straits; RW87-37; 21 May 1987; 4–8m. USMN
341618; (2, 46–56); Palawan Is., Bararin I., SP 78-20; 23 May 1978. USNM 341626; (1, 51); Pamilikan I.; SP
78-43; 12 June 1978. USNM 171386 (A23294-295); (2, 57–58); Tataan, Simulac I; 19 Feb 1908. USNM
112161 (A.23555); (1, 55); USNM 171397 (A23868); (1, 38); Rapu Rapu I.; 22 June 1909; USNM 112132;
(A.23189, 23190, 23562, 23564) (11, 43–53); Romblon I.; 25 Mar. 1908; Albatross; x-ray. Sulu Sea: FMNH
22035; (1, 57); Jolo, Sulu Province; 20 July 1929; USNM 213154 (A23698); (1, 55); Jolo I; 6–7 Mar 1908;
USNM 171393 (A23516); (1, 44); Tulayan I, near Jolo I; 15 Sept 1909; USNM 171410 (A15585); (1, 49);
Tulayan I, near Jolo I; 15 Sept 1909; USNM 171089; (2, 48–49); Bubuan I.; 14 Feb 1908; Albatross Exp.;
USNM 112176 (A.23700); (1, 49); Jolo I; 6–7 Mar 1908; x-ray; USNM 112173 (A.23506); (1, 51); USNM
171395 (A.23505, 23507); (2, 48–51); Dalaganem I, off Palawan I; 8 Apr 1909; USMN 169723; (4, 43–54);
Zootaxa 1691 © 2008 Magnolia Press · 27
status of Nectamia & 4 new species
Machesi I., Palawan Is; Albatross; 5 Apr 1909; x-ray; USNM 112170 (A.23228); (1, 50); Palawan I; 28 Dec
1908; Philippine Sea: USNM 171385 (A23286); (1, 53); Biri Channel; 1 June 1909. USNM 112238
(A21863, 23814); (2, 62–65); Biri Channel; 1–2 June 1909. Samar I., Port Palapag: USNM 112240
(A.23273); (1, 50); 2 June 1909; USNM 176164; (7, 51–57); Albatross; x-ray; USNM 112131 (A.15337); (1,
54); x-ray. USNM 171389 (A.23362-363); (2, 50–57); USNM 112182 (A.23361); (1, 61); 3 June 1909; x-ray.
Luzon I. Gubat Port; USNM171402 (A.11868); (1, 57); 23 June 1909. USNM 112239 (A11867); (1, 65); 22
June 1909. SU 21038 (Tin Tag 3882); (1, 61); Bacon I; C.J. Pierson coll. South China Sea: USNM 171413
(A.8667); (1, 52); Manila Harbor; 3 Jan 1909. USMN 171391 (A23365); (1, 51); USNM 171407 (A.16653);
(1, 55) Galera Bay, Mindoro I; 9 June 1908. USNM 112135; (A.16315, 23340, 23355, 23357) (4, 43–55);
Endeavor Str., NW Coast Palawan I.; 22 Dec. 1909. Sulawesi Sea: USNM 171399 (A.24005); (1, 56); Sitanki
Reef, S. Tawi Tawi Group; 24 Sept 1909. USNM 171409 (A.16586, 16588); (2, 45–52); Tulnalutan I, E of
Zamboanga; 9 Sept 1909. USNM 112179 (A.14055); (1, 48); USNM 171408 (A.8072, 15781-2) (3, 35–52);
USNM 171404 (A.14056); (1, 50); Sacol I. E of Zamboanga; 9 Sept 1909. USNM 112180 (A23475); (1, 53);
Tapiantana I: 13 Sept 1909. USNM 171401 (A24058); (1, 44); USNM 171411 (A.4642); (1, 47); Tonquil I; 14
Sept 1909. Camotes Sea: BPBM 36849 (1, 39); Mactan I.; 12 Oct.1995. Bohol Sea: USNM 171412 (A.5743,
5745); (2, 57–66); USNM 112183 (A.5744); (1, 54); Camiguin I; 3 Aug 1909. BPBM 28580; (2, 56–65);
Negros I., Dumaguete; 5m; 4 June 1981; Samar Sea: USNM 171382 (A.23991); (1, 45); Cataingan Bay,
Masbate I.; 18 Apr 1908. Sibuyan Sea: USNM 169727; (1, 49); Marinduque I; Albatross; 24 Apr 1908; x-ray.
Mindanao I.: USNM 171414 (A.9215); (1, 42); Mindanao; 6 Aug 1909. USNM 171384 (A.20570, 23217-
18); (3, 53–59); USNM 112174 (A.23215-16); (2, 59–63); Opol; 4 Aug 1909. Australia: Coral Sea: Lizard
I.: AMS P.24725; (2, 42–60), ROM 44265; (4, 37–70); Escape Reef; 6 Nov 1981; 2–11 m. AMS P.24743; (1,
48); One Tree I.: USNM 213158; (2, 67–68); 1 Dec 1966; VGS 66–14; USNM 213160; (1, 40); 11 Dec 1966;
VGS 66-19. MCZ 38496; (1, 59); Curacoa I; 28 Sept 1952. Papua New Guinea: USNM 341634; (17, 23–
65); Hermit I.; VGS 78-19; 4 Nov. 1978. BPBM 32581; (1, 55); Kranket Island; 1–3m.; 12 November 1987.
USNM 213162; (1, 37); Kuia I., Trobriand Is.; 11 June 1970; BBC-1514. USNM 213163; (8, 28–68); Kiri-
wina I.,Trobriand Is.; 8 June 1970; BBC-1509. Carolina Islands: Palau: BPBM 31401; (5, 16–55); Uleb-
sechel I.; 7–15 m; 9 Jul 1986. Urukthapel Island: CAS 84071; GVF 1979; (1, 56). Kayangel Island: CAS
83799; GVF 867; (37, 27–51); 26 Aug 1956. CAS 84083; GVF 866; (4, 35–49); 26 Aug 1956. Aulong Island:
CAS 84056; GVF 1991; (3, 46–56); 12 Oct 1956. CAS 84057; GVF 672; (1, 66); E of Malaklal Passage; 18
Sept 1955. CAS 83789; GVF 493; (1, 36) Auluptagel I.; 9 Oct 1957. Mariana Islands: Saipan: CAS 80792;
(2, 48–53); G90-8; 24 Feb 1990. Yap: CAS 83795; GVF 1924; (1, 54); 29 Dec 1959; Sta. 101. Kapingama-
rangi Atoll: CAS 83965; GVF 475; (19, 19–51); 20 Aug 1954; Sta. 172. CAS 83805; GVF 479; (19, 23–58);
21 Aug 1954; Sta. 176. Ifaluk Atoll: CAS 83793; GVF 136; (4, 37–49); 25 Sept 1953; Sta. 15. CAS 83879;
GVF 185; (1, 61); 8 Oct 1953; Sta. 64. Pohnpei: CAS 98023; GVF 493; (1, 40); 4 Oct 1954. Marshall
Islands: Bikini Atoll: FMNH 60028; (2, 26–28); Bikini I; 14 Aug 1946; S-46-349; USNM 142416; (1, 51);
off Amen I; 4 Aug 1946; LPS, Sta. S-46-307. Majuro Atoll: USNM 179404; (1, 51);Rita I; 12 Mar 1958;
Bartsch Sta. Solomon Islands: FMNH 22032-033; (2, 43–52); Ugi I; 10 Apr 1929; Crane Pacific Exp. BPBM
17375; (5, 33–69); black & white; Guadalcanal, Honiara; 16 m; 4 Aug. 1973. Santa Cruz Islands: USNM
213166; (4, 18–45); Vanikoro I; 3 Sept 1963; Te Vega Cr. 1, Sta. 35. USNM 213344; (14, 33–69); Vanikoro I;
16 Apr 1965; Te Vega Cr. 7, Sta. 259. New Caledonia: ROM 65766; (5, 64–69); 22°20'30"S 166°14'45"S;
RW91-19; 11 Sep 1991. USNM 324845; (2, 42–43); Noumea; 7 Nov 1991. Loyalty Islands: USNM 324847;
(1, 70); Ouvea Atoll; 17 Oct 1991. Tonga Island: USNM 213167; (121, 15–73); Nivatobutabu I; 31 May
1965; Te Vega Cr. 7, Sta. 301. USNM 213168; (80, 34–65); Neiafu, Vava’u; 28 June 1965; Te Vega Cr. 8, Sta.
308. Tongatapu Group; USNM 334689; (1, 56); JTW 93-4; 23 Oct. 1993. USNM 334704; (2, 51–64); 28 Oct
1993. USNM 334705; (5, 65–73); 27 Oct 1993. USNM 334427; (2, 57–82); JTW 93-8; 27 Oct.1993. Fiji:
CAS 223738; (2, 24–29); Muaivuso, 18°08.836'S, 178°21.988'E; G0-52; 11 Feb 2002; 11–14 m. CAS
223746; (3, 36–67); reef face in keyhole, 16°49.864'S, 178°19.625'E; G02-133; 3Apr 2002; 3–7 m. CAS
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28 · Zootaxa 1691 © 2008 Magnolia Press
223748; (4, 35–64); NE of Yaqaga I., 16°30.717'S, 178°38.730'E; G02-106; 25Mar 2002; 3–6 m. CAS
223749; (2, 44–44); Balavu Harbor, 17°11.212'S, 179°00.095'W; G03-24; 8 Jan 2003; 9–15 m. CAS 223750;
(6, 40–56); E of Nananu Passage, 17°15.063'S, 178°13.445'E; G02-80; 16 Mar 2002; 6–8 m. CAS 223751; (6,
39–50); Nasava Bay, 17°29.442'S, 178°21.171'E; G02-143; 4 Nov 2002; 5–8 m. CAS 223756; (8, 29–61); S
of Delavalau, Savuna Pt., 16°56.902'S, 178°58.231'E; G02-99; 21 Mar 2002; 1–4 m. CAS 223758; (9, 29–
53); S of Kia I., 16°16.911'S, 179°09.335'E; G02-121; 30 Mar 2002; 7 m. CAS 223759; (10, 25–50); Bua Bay
off Naicobocobo Pt. Reef; 16°50.139'S, 178°31.553'E; G02-102; 24 Mar 2002; 1–4 m. CAS 223760; (1, 52);
between Ovalua and Moturiki, 17°44.477'S, 178°45.334'E; G02-159; 8 Nov 2002; 1–9 m. USNM 213169;
(20, 29–80); Wailagi Lala I; 26 May 1965; Te Vega Cr. 7, Sta. 295. ANSP 128369; (7, 36–45); Suva; FIJ74-3;
15 Apr 1974. BPBM 11406; (2, 52–55);color, black & white; Joske Reef; 6 m; 3 Sep. 1971. CAS 223747; (1,
37); N side of Viti Levu, 17°16.704S, 178°12.932E; G02-186; 15 Nov 2002; 3–6 m. CAS 223736; (8, 38–52);
Davetia Bay; 16°45.578'S, 179°54.618'E; G03-66; 26 May 2003; 7–12 m. CAS 223752; (5, 30–36); Davetia
Bay, 16°45'.578'S, 179°54.618'E; G03-67; 26 May 2003; 1–3 m. CAS 223761; (16, 27–55); Davetia Bay,
16°45.578'S, 179°54.618'E; G03-66; 26 May 2003; 7–12 m. CAS 223762; (42, 26–55); Nasau Bay,
16°43.650'S, 179°53.970'E; G03-62; 25 May 2003; 1–4 m. CAS 223753; (4, 30–48); Nasau Bay, 16°43.650'S,
179°53.970'E; G03-61; 25 May 2003;0–3 m. ROM 44178; (4, 21–28); off Suva; 12 Apr 1982. ROM 43262;
(51, 12–70); Great Astrolabe Reef; WE 83-21; 27 Mar 1983. ROM 43263; (2, 10–62); Great Astrolabe Reef;
WE 83-9; 20 Mar 1983. USNM 212841; (4, 34–45); Mbulia I., 18°50'30"S 178°32'10"E; 28 Aug 1963; 3–6m.
USNM 213170; (9, 41–57); N. of Vuro I, Great Astrolabe Reef; 8 May 1965; Te Vega Cr. 7, Sta. 278. USNM
213343; (23, 19–59); N. of Vuro I, Great Astrolabe Reef; 8 May 1965; Te Vega Cr. 7, Sta. 279. Wallis Is.:
USNM 369974; (80, 23–56); Ila Uvea, 13°19'17"S 176°15'10"W; 14 Nov 2000. Samoan Islands: Western
Samoa: Upolu: FMNH 4804; (3, 50–60); Apia; 1902; D.S. Jordan coll. IUM 10894; (2, 40–62); Apia; 1902.
USNM 52432; (18, 46–68); Apia, one spec. drawn. USNM 126257; (2, 65–73); Apia; 1902; US Fish Comm.
MCZ 30268; (1, 64); Apia; 1902, No. 06556. American Samoa: Tutuila: SU 8888; (13, 58–71) Pago Pago
1902. USNM 58522; (8, 59–63); Pago Pago; 1902; US Fish Comm. Phoenix Island: USNM 213171; (32,
30–69); Kanton I; 12 July 1965; Te Vega Cr. 8, Sta. 314. Cook Islands: FMNH 16239-40; (2, 44–50); Aitut-
aki; 1930; Chancellor-Field Exp. USNM 213172; (10, 13–30); Tongareva (Penrhyn) Atoll, S. of Molokai I; 13
June 1965; S.I. POBS coll. Society Islands: Tahiti: FMNH 22004-06; (3, 37–62); Maraa; 18 Feb 1929; Crane
Pacific Exp. USNM 164557; (4, 31–64); lagoon. Mangareva MCZ 29552; (10, 64–74).
Diagnosis. A Nectamia with no dark saddle on body below the second dorsal fin, many pale bars on body,
caudal peduncle with an incomplete dark wide bar, mostly above the lateral line, caudal fin with dark margins,
subocular mark, thin and triangular (Tab. 2, Fig. 11); 13 pectoral-fin rays; total gill rakers usually 25–31 (Tab.
7); body depth 39–44%, caudal-peduncle depth 16–20%, second anal-spine length 17–21%, and pectoral-fin
length 26–29% of standard length (Tab. 3).
Description. For general body shape see Figure 11. Range of proportions (as percentage of standard
length) in Table 3 with paratypes and non-type material in parentheses.
Dorsal fin VII(I)–I,9; anal fin II,8; pectoral fin usually 13-13; pelvic fin I,5; principal caudal rays 9+8;
pored lateral-line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line
5–6; median predorsal scales 3; circumpeduncular scale rows 12 as 5+2+5; total gill rakers 28 (25–31), well
developed 25 (23–29), upper arch 2+5 (0-2+5-7), lower arch 20+1 (18-20+0-1), second arch with shorter gill
rakers 2+16.
Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and
vomer; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10+14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural. Three
supraneurals, two supernumerary spines on first dorsal pterygiophore. Basisphenoid present. Supramaxilla
absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal margins.
Infraorbital shelf present on third bone.
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status of Nectamia & 4 new species
Holotype with ctenoid scales on cheek, subopercle, opercle, nape, isthmus, cycloid at base of pectoral fin,
ctenoid on rest of body, base of pelvic with missing scales, large scale missing exposing a small cycloid axil-
lary scale. Pored lateral-line scales complete, fourth scale simple with a single pore above with a large anterior
and a small central foramen and below main canal one exposed and one covered pore by a body scale (Fig.
4E). Base of pelvic fin with two large weakly ctenoid scales in paratypes and other material.
Etymology. The Latin word luxuria, meaning profusion, referring to the numerous pale bars. Treated as a
noun in apposition.
FIGURE 11. Nectamia luxuria, holotype, USNM 209644, 59.1 mm SL, Indonesia, Ceram, Tandjung Liang, 10 Jan
1973, 0–7 m, Rumphius Exp. I, Sta. Li-1; VGS 73-6.
TABLE 7. Geographic variation in gill-raker and rudiments counts for Nectamia luxuria.
* = holotype
Upper Arch Lower Arch Total
678x 18 19 20 21 22 23 x 25 26 27 28 29 30 31 x
INDIAN OCEANMaldive Is 10 5 7.3 5 10 20.7 3 9 3 28.0
Cocos-Keeling Is 2 8.0 1 1 22.5 1 1 30.5
Pulo Bai, Indonesia 1 13 7 7.3 6 15 20.7 5 11 5 28.0
PACIFIC OCEAN
Hong Kong 5 97.6 4541 20.1 15521 27.8
Vietnam 2 17.3 111 20.0 11 1 27.3
Thailand 6 2 7.3 5 2 1 20.5 3 4 1 27.8
Malaysia 1 3 1 7.0 4 1 21.2 1 3 1 28.2
Philippines 5 18 4 7.0 4 12 10 1 19.3 6 8 10 1 1 26.3
Sabah, Darvel Bay 15 7.0 1491 19.7 1491 26.7
Indonesia 2 18* 3 7.0 6 14 3* 19.9 8 10 3* 2 27.0
Palau / Saipan 9 2 7.2 1 3 7 20.5 3 6 2 26.9
Ifalik / Pohnpei 4 1 7.2 2 2 1 20.8 2 1 2 28.0
Coral Sea 2 77.8 153 21.2 333 29.0
New Caledonia 1 1 7.5 1 1 21.5 1 1 29.5
Bismark Archipelago 5 1 7.2 4 2 20.3 4 1 1 27.5
Fiji 4 67.6 361 20.8 1531 28.4
Wallis I. 1 4 7.8 5 21.0 1 4 28.8
Tonga Is 3 8.0 1 2 21.7 1 2 29.7
Samoa Is11167.3 1197 21.2 11844 28.5
SUMMARY
Maldives / Sumatra 1 23 14 7.3 11 25 1 1 20.8 8 20 8 1 1 28.1
China Sea 1 16 13 7.4 5 11 11 3 20.4 2 10 12 4 2 27.8
Philippines / Indonesia 7 51 7 7.0 5 22 33 5 19.6 7 20 29 5 3 26.6
Palua / Pohnpei 1337.2 1591 20.6 3832 27.3
Coral Sea / Samoa 1 18 29 7.6 1 3 27 17 21.3 1 1 15 17 14 28.9
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30 · Zootaxa 1691 © 2008 Magnolia Press
FIGURE 12. The distribution of collection sites for Nectamia luxuria with regional total gill raker averages.
Zootaxa 1691 © 2008 Magnolia Press · 31
status of Nectamia & 4 new species
Life colors. See Figure 6B. Head, body and fins without stripes; head with dark oblique cheek mark from
eye to angle of preopercle, lachrymal and anterior infraorbital with yellowish mark, iris brownish with white
inner ring; head and body bronze to brownish background with alternating pale and brown bars, center of
brown bars commonly with narrow light region; caudal peduncle with a dark basicaudal mark mostly above
pored lateral-line scales, fading or absent below pored lateral-line scales, partial pale region just anterior to
peduncle mark; first dorsal fin with dusky membranes between spines II–IV, second dorsal, pectoral, pelvic
anal and caudal fins pale except for yellowish edges to the second dorsal, anal and caudal fins, base of pecto-
ral fin yellowish.
Preserved color pattern. Holotype: Head with darkish mark on lachrymal and anterior infraorbitals,
well-defined dark angular cheek tapering to a point at preopercular ridge, rest of head without markings; body
with alternating brownish and lighter bars from opercle flap to near posterior origin of second dorsal fin,
darker area below second dorsal fin, caudal peduncle with dark bar above pored lateral-line scales; first dorsal
fin dusky, pectoral fins pale, second dorsal fin, anal fin and pelvic fins all dusky to pale, caudal fin darkish on
several of outer rays forming dark edges, interior of caudal fin pale (Fig. 11); stomach and intestine blackish;
roof of mouth and upper gill arches pale. Adults (Tab.2): wide, triangular subocular cheek mark behind upper
jaw; first dorsal fin with dark membrane between spines 1–4; edges of caudal lobes dark; caudal peduncle
with a incomplete dark, bar mostly above the lateral-line; no dark saddles on body; many alternating bars on
side of body (7–13); stomach and intestine black; roof of mouth mostly pale with scattered tiny melanophores
not extending onto gill arches. Juveniles: similar to the adults except the caudal mark extends below the lateral
line and usually fewer bars.
Distribution. Known from the Maldive Islands eastward in the Indian Ocean and in the West Pacific to
Mangareva Island (Fig. 12). This species is usually taken in depths less than 11 m.
Remarks. Jordan and Seale (1906:240, fig. 33) illustrated this species from Apia, Western Samoa as
Apogon savayensis. Kuiter and Kozawa (1999, p. 34, A–D) provide four color photographs as Apogon sp 20.
None of the other species of Nectamia has as many pale body bars. Weber and de Beaufort (1929) listed Jor-
dan and Seale’s A. savayensis as a questionable name in the synonymy of Apogon bandanensis.
Average gill-raker counts differ by 1–2.5 elements with geography (Fig. 12, Tab. 7). The broad range of
gill-raker elements is a polymorphic character. The lowest average was found in the Philippino-Indonesian
region. There were no obvious differences in preserved markings or in the available underwater photographs
to suggest recognizing additional taxa. Examination of DNA variation may bring a finer focus to populations
of Nectamia luxuria.
Nectamia savayensis has a similar dark caudal-peduncle mark above the lateral-line, colored edges on the
caudal fin lobes but just a few variably faint pale body bars and a broad cheek mark. Nectamia luxuria has
many pale bars on the body in life and preservation, yellow edges to the caudal fin in life, that become dark in
preserved material and a narrow cheek mark. Robert Myers, personal communication, has not seen this spe-
cies in Guam and I found no material from Guam which could be conclusively be identified as this species. It
should occur in Guam.
This species has been taken at stations with one or more of the following species: Nectamia fusca (CAS
28511, 84087, 84103, 95967, 95984, FMNH 22031, 22036, SU 26650), N. savayensis (ANSP 12245, CAS
83800, USNM 112047, 261147, 261148, 261158), N. similis (USMN 112130, 112134, 112136, 171387,
213610, 261024) and N. viria (ANSP 77366, USNM 153237, 210960, 213111, 213127, 13128, 213341,
268256, 268271, 319149, 341619, 341627). Specimens (51–58 mm SL) in CAS 223761, SU 51220, ROM
54318 and USNM 171386 showed buccal enlargement, but no eggs were found. The 54 mm SL specimen in
USNM 112183 had eggs in its mouth. The largest of the 1491 specimens was 77 mm SL.
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32 · Zootaxa 1691 © 2008 Magnolia Press
Nectamia savayensis (Günther, 1872)
Figures 6F, 13, 18 and Tables 2–3, 8
Type material: Holotype Apogon savayensis BMNH 1871.9.13.218; 48 mm SL; Savai’i (Savay, Western
Samoa); Godeffroy Coll.
Additional Material: Indian Ocean: Mozambique: SAIAB 3311; (2, 47, 53); Island das Rolas; 11 Aug
1951; x-ray. SAIAB 3318; (1, 58); Pinda Reef; 9 Sept. 1956; x-ray. SAIAB 3321; (1, 42); Mozambique, Pinda
Reef, June 1951; x-ray. SAIAB 3315; (2, 36–55); Pinda Reef; July 1950; x-ray. SAIAB 3317; (1, 65); Matemo
I.; 13 Aug 1951; x-ray. Zanzibar: SAIAB 3320; (1, 56); 3 Sept 1952; x-ray. Madagascar; UMMZ 185908;
(1, 44); Tulear; 11–13 July 1964; JB Sta. 64-22. Aldabra Islands: USNM 213130; (5, 31–49) Lagoon inside
SE portion of West I (Ile Picard); 3 Dec. 1964; IIOE Cr. 9, HAF-16. SAIAB 3314; (1, 68); Nov 1954; x-ray.
Farquhar Islands: USNM 213131; (21, 50–67) 6 Dec 1964; IIOE Cr 9, R. Suttkus-37; USNM 213132; (74,
24–62) Lagoon W. of North I; 6 Dec 1964; IIOE Cr. 9; HAF-18. Comoro Islands: USNM 213133; (6, 56–68)
Grande Comore I; W. side edge of fringing reef; 27 Nov 1964; IIOE Cr 9, HAF-14. USNM 213134; (23, 27–
60); Lagoon ca 3.5 mi. SW of S Pt of Pamanzi I.; 24 Nov 1964; IIOE Cr. 9; HAF-9. Amirante Isles: ANSP
152769; (4, 39–71) D’Arros I; 5 Mar 1964; Sta. F-88. USNM 213135; (50, 25–67) St. Joseph Is, Ressource I;
8 Dec 1964; IIOE Cr 9, H.A. Fehlmann-19 ANSP 152792; (1, 14) D’Arros I; 5 Mar 1964; Sta. F-87. ANSP
164655; (1, 14); off N. end of North I; 3 Mar. 1964; IIOE-ANSP; Sta. F-78. USNM 213136; (1, 17); D’Arros
I; 8 Dec 1964; IIOE Cr 9, Suttkus-40. Seychelles: Mahé I.: ANSP 152775; (2, 32–39); 19 Mar 1964; Sta. F-
119. ANSP 152791; (7, 41–67) Beau Vallon Bay; 15 Mar 1964. ANSP 152773; (6, 36–70); North West Bay; 9
Feb 1964; Sta. F-36. ANSP 152796; (16, 29–74); 31 Jan –2 Feb 1964; Sta. F-13. ANSP 152800; (5, 22–36)
Souris I.; 16 Feb 1964; Sta. F-53. CAS 58762; (2, 68–73); Te Vega; 27 Oct 1964. SAIAB 3322; (1, 57); 16
Sept 1954. SAIAB 3319; (2, 64–65); 27 Sept 1954. BPBM 35518; (1, 19); 24 Dec 1992. Beacon I.: ANSP
152801; (3, 29–45); 25 Jan 1964; IIOE-ANSP; Sta. F-5. ANSP 152802; (2, 28–39); 3 Feb 1964; Sta. F-18.
ANSP 152772; (1, 18); Off Port Victoria I.; 6 Feb 1964; Sta. F-26. ANSP 152799; (2, 35–69); 27 Jan 1964;
Sta. F-7. ANSP 152798; (5, 30–73); N. of Anonyme I, Mahé vicinity; 10 Feb 1964; Sta. F-37. ANSP 152797;
(5, 22–62) W. of NW Anonyme I., near Mahé I; 11 Feb 1964; Sta. F-44. Sta. F-114. ANSP 152771; (2, 24–39)
Just N. Anonyme I., near Mahé I; 2 Feb 1969; Sta. F-17. ANSP 152805; (14, 27–63) Between Praslin &
Round Is.; 22 Feb 1964; Sta. F-59. ANSP 152804; (1, 38) S of Round I, near Praslin I; 22 Feb 1964; Sta. F-61.
ANSP 152458; (1, 14); Curieuse Island; F-64 ; 23 Feb 1964. ANSP 152770; (11, 34–40) SE Rouge Pt,
Curieuse I; 23 Feb 1964; Sta. F-62. ANSP 152803; (6, 35–60); S. of Rouge Pt, Curieuse I.; 24 Feb 1964; Sta.
F-66. ANSP 152774; (9, 20–65) Faon I; 24 Jan. 1954; Sta. F-3. CAS 35363; (23, 53–67); Cerf I.; Te Vega Cr.
5; 21 Oct 1964. CAS 35369; (15, 55–76); St. Anne I.; Te Vega Cr.5; 19 Oct 1964. SAIAB 3312; (1, 58) Sil-
houette I; 2 Oct 1954; x-ray. Reunion: USNM 345778; (1, 34); west coast near St Leu; 7 Feb 1997. Mauri-
tius: FMNH 75711; (57, 42–74); Black Rock Point; 24 May 1964; LPW-41. USNM 349771; (11, 35–78);
M28; 16 May 1995. USNM 213137; (20. 43–77); reef off Port Louis; 12 June 1964; IIOE Cr. 6. USNM
213138; (1, 73); reef off Port Louis; 11 June 1964; IIOE Cr. 6. SAIAB 2120; (4, 40–92); La Morne Brabant;
THF-SA-28;7 Mar 1971; x-ray. SAIAB 2311; (11, 33–38); Palamar; THF-SA-44; 27 Mar. 1971; x-ray.
SAIAB 2234; (9, 44–52); near Riambel; THF-SA-42; 26 Mar. 1971; x-ray. BPBM 20102; (3, 46–67);Trou
d’Eau Douce; 31 Oct. 1973. St. Brandon Shoals: SAIAB 1893; (3, 66–71); Tortue I; 15 Mar 1971; THF SA-
35. SAIAB 2207; (1, 44); Raphael I.; 18 Mar 1971; THF SA-36. Chagos Archipelago: Diego Garcia Atoll:
USNM 213139; (2, 26– 55); 7 July 1967; HAF 67-36. USNM 213140; (2, 31–62); 26 June 1967; HAF 67-18.
USNM 213141; (18, 26–59); 13 June 1967; HAF 67-5. USNM 213142; (10, 30–54); 12 June 1967; HAF 67-
3. USNM 213143; (2, 32–67); 9 July 1967; HAF 67-38. USNM 213144; (3, 29–37); 20 July 1967; HAF 67-
49. USNM 213145; (2, 47–52); 18 June 1967; HAF 67-11. Maldive Islands: FMNH 75707; (14, 22–38)
Addu Atoll; 10 May 1964; LPW-40. FMNH 75670; (4, 32–52) Tiladummati Atoll; 31 March 1964; LPW-29.
FMNH 75683; (2, 41–44) Ari Atoll; 22 Apr 1964; LPW-35. FMNH 75652; (2, 66–67) S. Malé Atoll; 18 Mar
Zootaxa 1691 © 2008 Magnolia Press · 33
status of Nectamia & 4 new species
1964; LPW-24. FMNH 75695 (19, 22–59) S. Nilandu Atoll; 24 Apr 1964; LPW-37. FMNH 75665; (77, 25–
62) Miladumadulu; 27 Mar. 1964; LPW-28. FMNH 75698; (115, 21–65); Addu Atoll 30 Apr 1964; LPW-38.
BPBM 33049; (1, 62); North Malé Atoll, W. side of Huraa Island; depth 20–22m.; 24 Mar. 1988.SMF 4750;
(7, 28–40); Ari Atoll, 3°15´N, 73°00´E; 17 Feb 1958. SMF 4761; (2, 43–46); Fadiffulu Atoll, 3°15´N,
73°00´E; 13 Apr 1958. Sri Lanka: USNM 213146; (1, 59); Trincomalee; 4 Apr 1970; CCK 69-136. USNM
213147; (1, 42); Hikkaduwa, 1¼ mi. off H Kralla; 17 Feb 1970; CCK 69-109. USNM 213148; (1, 64); Korali
Pattu, Batticaloa District; 6 June 1970; TI 70-344. USNM 213149; (1, 15); Trincomalee; 8 Apr 1970; CCK
69-143. USNM 213150; (1, 22); Trincomalee; 27 June 1969; WSV 69-120. India: FMNH 75638; (3, 20–29);
Mandapan Camp; 31 Jan 1964; LPW-15. FMNH 75627; (3, 68–82); Gulf of Mannar, Musal tivu; 27 Jan 1964;
LPW-13. FMNH 75616; (9, 28–73); Gulf of Mannar, Musal tivu; 7 Jan 1964; L.P. Woods - 5. FMNH 75605;
(9, 62–66); Gulf of Mannar, Musal tivu; 30 Dec 1963; LPW-3. Nicobar Is.: SMF 7759; (2, 38–54); Tillan-
chong I., 8°00'N ,93°30'E; 3 Sep 1958. Cocos-Keeling Islands: ANSP 128377; (1, 22); North Keeling I; 6
Mar 1974; Sta. 22. ANSP 128374; (4, 20–22) West I; 28 Feb 1974; Sta. 12. ANSP 128372; (1, 30); West I; 25
Feb 1974; Sta. 6. ANSP 128376; (1, 27); Prison I; 7 Mar 1974; Sta. 23. ANSP 128373; (1, 16–17);Direction
Island; Sta 9; 27 Feb 1974. ANSP 128375; (1, 15–16);Horsburgh Island; Sta 5; 25 Feb 1974. Thailand: ROM
70484; 12(16–89); Phuket, 7°55'15"N 98°16'16"E; RW 93-8; 13 Nov 1993; 8–13 m. ROM 71478; 40(44–74);
Phuket, 7°47'10"N 98°23'45"E; RW 93-3; 10 Nov 1993; 0–9 m. ROM 71479; 20(33–66); Phuket, 7°45'40"N
98°20'2"E; RW93-4; 10 Nov 1993; 0–6 m. ROM 72174; 4(12–58); Phuket, 7°73'15"N 98°22'50"; RW 93-12;
16 Nov 1993; 3–8 m. ROM 72176; 1(54); Phuket, 7°72'15"N 98°19'35"E; RW 93-17; 18 Nov 1993; 15–22 m.
ROM 72177; 9(38–62); Phuket, 7°72'15"N 98°19'35"E; RW 93-18; 18 Nov 1993; 1–9 m. ROM 72179;
1(62.5); Phuket, 8°83'99"N 97°39'99"E; RW 93-34; 14 Dec 1993; 0–10 m. USNM 213152; (1, 33); Paton
Bay, Phuket; 22 Mar 1963; IIOE Cr. I. Indonesia: CAS 95833; (1, 57); Pulau Pandan, off Padang, Sumatra;
25–26 m; 17 Apr 1997. Australia: Timor Sea: CAS 52558; (29, 24–45); Ashmore I.; 3 Jan. 1973. Pacific
Ocean: Thailand: CAS 213345; GVF 2172; (1, 53); Khorn Aho Bay, 12 37'37"N 101 20'31"E; 24 Apr 1960.
CAS 84058; GVF 2651; (1, 72); 1.5 mi. ESE of Town of Prachuap Khiri Khan; 19 June 1961. Viet Nam: CAS
83817; GVF 2113; (10, 59–70); Ilot du Sud, 10°29'15"N; 108°57'30"E; 10 Mar 1960; MV Stranger, Sta. 60-
96. ROM 79172; (6, 19–79); RW02-21; 9 Jun 2003 Singapore: BPBM 21854; (1, 50); Sulu I.; 24 July 1977.
Indonesia: CAS 62476; (1, 43); Sangi Siang I.; JEM 87-7; 24 Sep 1987. Java Sea: BPBM 18583; (1, 49);
Capelan Seribu, Pulau Petri; black and white; 17 Feb. 1975. Banda Sea: USNM 213113; (8, 43–73); SE Naira
I, Banda I.; 8 Mar 1974. VGS 74-9; USNM 213187; (10, 32–69); Ambon; 14 Mar 1974; VGS 74-13. USNM
171388 (A.23359); (1, 47); Bauru I; 9 Dec. 1909; Molucca Sea: Sulawesi: USNM 112234 (A.23394); (1, 63)
Gulf of Tomiki; 16 Nov 1909. USNM 112190 (A.23974); (1, 49) Limbs Str.; 11 Nov 1909. Sulawesi Sea:
BPBM 36709; (1, 67); Sulawesi, Maenad, Bunaken; 8m; 28 Oct 1991. Philippines: USNM 344522; (15, 32–
69); Panay, Lawigan, Tiolas; JTW 95-4. USNM 261148; (9, 48–60); Negros I., south tip; SP 78-9; 12 May
1978. USMN 261151; (26, 21–62); Palawan Is., Bararin I., SP 78-20; 23 May 1978. USNM 261147; (2, 65–
71); Pamilikan I.; SP 78-43; 12 June 1978. USNM 329215; (38, 28–62); Negros Oriental, Siquijor I.; LK 79-
15, 15 May 1979. Sulawesi Sea: USNM 171399 (A.24005); (1, 56); Sitanki Reef, S. Tawi Tawi Group; 24
Sept 1909; Sulu Sea: ANSP 98137; (15, 34–52); Sibutu Island; 27 Apr 1926. USNM 171403 (A.21558); (1,
52); Pilas I, S of Zamboanga; 12 Sept 1909; USNM 171405 (A.15565); (1, 40); Bugsuk I, N. Balabac Str.; 5
Jan 1909. Philippine Sea: USNM 112164 (A.23655); (1, 54); Batan I; 8 June 1909; x-ray. China: CAS
83800; GVF 1750; (8, 51–64); Pratas Reef; 25 May 1958; Sta. HK 73. Taiwan: USNM 213155; (4, 49–60);
Bay between K’en-ting and Ta-yuan Shan; 1 May 1968; VGS 68-12. USNM 213156; (3, 65–72); Off Ch’uan-
fan-shih, SW shore; 3 May 1968; VGS 68-14. USNM 213157; (13, 44–82); Ch’uan-fan-shih; 23 Apr 1968;
VGS 68-2. BPBM 23300; (4, 37–83); S end at Ch’uan-fan-shih; 0–6m; 16 Jul 1978. South Korea:
CAS29963; (1, 18); Inchon area;6–7 Jun 1973. Carolina Islands: Palau: BPBM 7225; (5, 37–54); SE of
Auluptagel I.; 26 Sep. 1966. BPBM 8091; (6, 35–55); SW of Koror I.; 12 Jun 1968. CAS 83820; GVF 908; (1,
52); Ngadarak Reef; 25 Sept. 1956, Sta. 122. CAS 84055; GVF 925; (7, 29–61); Ngaruangl Reef; 5 Oct 1958;
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Sta. 139. CAS 83778; GVF 804; (3, 50–68); Ngadarak Reef; 12 Jul 1956. CAS 84061; GVF 897; (1, 74);
Ngadarak Reef; 12 Sep 1956. CAS 84085; GVF 1385, (3, 54–64); Ngadarak Reef; 11 Sep 1957. CAS 83790;
GVF 490, (2, 64–68); Urukthapel I.; 23 Sep 1954. CAS 84084; GVF 1397; (27, 17–61); Babelthuap I.; 22 Sep
1957. CAS 83791; GVF 1958; (3, 53–67); Ngethil I.; 13 Jul 1959. Ifaluk Atoll: CAS 83794; GVF 129; (22,
32–62); 21 Sept 1953; Sta. 8. CAS 83809; GVF 216; (17, 37–54); 20 Oct 1953; Sta. 95. CAS 83810; GVF
262; (21, 41–54); 28 Oct 1953; Sta. 141. CAS 83777; GVF 976; (10, 38–61); Ulitki Atoll; Sorlen I, W. side;
22 Sep 1956. CAS 83880; GVF 215; (13, 39–44); 20 Oct 1953; Sta. 94. Truk: BPBM 9313; (13, 14–56); 10
Jul 1969l. Pohnpei: CAS 83788; GVF 496; (13, 23–53); 6 Oct. 1954; Sta. 193. CAS 83784; GVF 493; (1,
72); 4 Oct 1954; Sta. 190; Kapingamarangi Atoll: CAS 83798; GVF 317; (3, 31–42); Tokoteihi; 1 July 1954;
Sta. 14. CAS 83970; GVF 329; (1, 45); 8 July 1954, Sta. 26. Sorol Atoll: CAS 83781; GVF 993; (1, 45); 28
Oct 1956. Mariana Islands: Guam: ANSP 85880; (1, 82); AS00-G-6-1; 1 Jun 1900. CAS 84065; GVF 788;
(25, 51–65); 14 Jun 1956. CAS 83877; (3, 40–47); GVF 1839; 10 Oct 1958; sta 11. CAS 213343; GVF 788;
(1, 62.9); 14 Jun 1956. UGM 1418; (3, 53–69); 5 Oct 1968; GFW-111. UGM 1415; 7(45–71); 28 June 1968;
GFW-119; 27–33m. Saipan ANSP 164649; (1, 66); SAI-1-25; 18 Oct 1953. CAS 83894; GVF 790; (4, 36–
63); 17 June 1956; Sta. 4. CAS 83896; (9, 16–35);GVF 789;16 Jun 1956. CAS 213326; (1, 56); G90-8; 24 Feb
1990. CAS 80770; (3, 47–51); G90-6; 25 Feb 1990. Yap I.: CAS 84155; GVF 1935; (1, 53); 6 Jan. 1959; Sta.
112. Marshall Islands: Arno Atoll: USNM 166599; (1, 29); Eonëb-je I.; 3 July 1950. Bikini Atoll: USNM
142415; (9, 19–32); Ari I.; 7 Aug 1946; LPS, Sta. S-46-308. Enewetak Atoll: BPBM 28971; (1, 51);
Enewetak Atoll; 23m; 24–25 Sep. 1982; x-ray. BPBM 31233; (1, 51); 19 Apr 1976. Kwajalein Atoll: USNM
142428; (1, 19); near Ennylabegan I.; 1 Sept 1946; LPS, Sta. S-46-397. Rongerik Atoll: USNM 142417; (2,
27–28); Latoback I.; 14 Aug 1947; LPS, Sta. S-1041.6. Rongelap Atoll: USNM 142414; (27, 23–61);
Kieshiechi I; 24 July 1946; LPS, S-46-285. Wake I.: BPBM12041; (1, 49); May 1957. SU 50057; (2, 47–48).
SU 51220; (24, 39–51); Between Wilkes and Peale Is; 20 Sept 1957. Papua New Guinea: Bougainville I.:
USNM 268266; (79, 31–67); TeVega Cr. 6; 19 Oct. 1984. USNM 262489; (9, 17–54); Hermit Is., Amot I.;
VGS 78-10; 30 Oct. 1978. USNM 213251; (2, 19–31); Kranket I.; BBC-1498; 31 May 1970. USNM 296755
(1, 47) and CAS 65301 (1, 44); Wongat I.; SGP 87-29; 20 May 1987. CAS 65299; (1, 54); Wongat I.; SGP 87-
33; 22 May 1987. Nagada Harbour: USNM 296845; (1, 38) and CAS 65308 (2, 44–52); SGP 87-23; 16 May
1987. USNM 296742; (1, 48) and CAS 65300; (1, 34–50); SGP 87-30; 20 May 1987. CAS 65302; (1, 44);
SGP 87-19; 12 May 1987. CAS 65305; (1, 46); SGP 87-17; 8 Apr 1987. CAS 65303; (3, 55–63); n. of Badilu;
SGP 87-27; 18 May 1987. CAS 65304; (1, 42); Tab (Pig) I.; SGP 87-21; 15 May 1987. USNM 213161; (5,
23–44); Madang Harbor; 26 May 1970; BBC-1488 and 1490; USNM 213164; (4, 47–58); Madang Harbor; 31
May 1970; BBC-1497.Australia: Coral Sea: ANSP 123307; (2, 27–29); Big Hope Island; TSA-33; 27 Jan
1969. ANSP 123325; (9, 25–30) Big Hope Island; TSA-22; 19 Jan 1969. ANSP 123415; (2, 10–18); Little
Hope Island; TSA-1; 3 Jan 1969. ANSP 123373; (1, 25); Northern Escape Reef; TSA-30; 24 Jan 1969. ANSP
122943; (3, 15–29); W of Little Hope Island; TSA-32; 26 Jan 1969. ANSP 174722; (2, 24–26); Big Hope
Island; TSA-34; 27 Jan 1969. ANSP 123347; (2, 22–25); Endeavour Reef; TSA-6; 6 Jan 1969. ANSP 123316;
(3, 38–67); Big Hope Island; TSA-21; 18 Jan 1969. ANSP 123401; (5, 22–25); TSA-5; 6 Jan 1969. ANSP
122215; (1, 65); Palm Group, Curacoa Is.; 28 Sep 1952. ANSP 135333; (3, 56–68); Yonge Reef; LZ-75-56;
23 Nov 1975. CAS 40118; (2, 23–60); 14 35' N 145 36' E, Yonge Reef; 23 Nov.1975. CAS 28509; (4, 28–
37); 14 0.78' S. 143 59.0' E, Wharton Reef; 9 Oct 1960; SIO61-132. USNM 218082; (3, 66–71); Yonge
Reef; 23 Nov. 1975. CAS 40118; (2, 23–59); Young Reef, 14 35'S 145 36'E; 23 Nov 1975; 7–15m. Solomon
Islands: USNM 213165; (6, 31–54); 10 Mar 1965; Te Vega Cr. 6, Sta. 245. USNM 357421; (31, 52–77);
Santa Cruz Is., 10°16'30"S 166°16'30"E; 18 Sep 1998; 0–13 m. Loyalty Islands: USNM 324847; (1, 70);
Ouvea Atoll; 17 Oct. 1991. New Caledonia: USNM 324845; (2, 43–43); Noumea, Pointe Cluxel; 7 Nov.
1991. Tonga Islands: USNM 341604; (7, 37–54); JTW 93-22; 5 Nov. 1993. USNM 341605; (4, 45–61); JTW
93-17; 3 Nov. 1993. USNM 341606; (5, 52–70); JTW 93-29; 11 Nov. 1993. USNM 341607; (1, 44); JTW 93-
42; 17 Nov. 1993. USNM 341608; (4, 33–36); JTW 93-11; 28 Oct. 1993. USNM 341609; (1, 35); JTW 93-15;
Zootaxa 1691 © 2008 Magnolia Press · 35
status of Nectamia & 4 new species
2Nov. 1993. Tongatapu Group: USNM 334704; (2, 51–64); N. Of Atata I.; 28 Oct 1993. USNM 334705; (5,
65–73); Malinoa I.; 27 Oct 1993. USNM 341628; (6, 40–78); JTW 93-8; 27 Oct. 1993. Fiji: CAS 83888;
GVF 2008; (1, 60) S. Yasarwas Is.; 4 Aug 1959. CAS 224788; (1, 38); Muaivuso, 18°08.836'S, 178°21.988'E;
G02-52; 11 Feb 2002; 11–15 m. CAS 224789; (5, 37–54); S of Kia I., 16°16.911'S, 179°09.335'E; G02-121;
30 Mar 2002; 6 m. CAS 223757; (2, 53–59); E of Nananu Passage, 17°15.063'S, 178°13.445'E; G02-82; 16
Mar 2002; 1–5 m. USNM 242056; (1, 49); Viwa I.; VGS 82-31; 27 May 1982. USNM 262432; (7, 41–70);
Naviti I.; VGS 82-32; 28 May 1982. USNM 288131; (3, 20–48); Rotuma; VGS 86-9; 14 May 1986. USNM
288195; (7, 28–65); Rotuma; VGS 86-6; 12 May 1986. USNM 262617; (22, 25–68); Lua Is., Yangasm Clus-
ter; VGS 82-16; 2 May 1982. USNM 262455; (29, 39–73); Totoya I.;VGS 82-9 27 April 1982. Samoan
Islands: American Samoa: BPBM 27929; (6, 62–75); Tutuila; 0.5–6m.; 2 Mar. 1972. USNM 58522; (1,
60.1) Tutuila; cleared & stained. Kiribati: Onotoa: USNM 167318; (9, 31–60). USNM 167319; (3, 36–63).
BPBM 15207; (3, 40–57); 1951. Society Islands: Bora-Bora: CAS 83882; GVF 1364; (1, 56); 10 Aug 1957;
Sta. 32. CAS 83883; GVF 1367; (2, 29–46); 14 Aug 1957; Sta. 35. CAS 83876; GVF 1366; (1, 53); 12 Aug
1957; Sta. 34. CAS 83886 GVF 1369; (2, 51–55); 16 Aug 1957; Sta. 37. SU 24737; (1, 59); 26 Feb 1929;
Huahine: CAS 83972; GVF 1358; (6, 34–44); 2 Aug 1957; Sta. 26. CAS 83889; GVF 1357; (1, 42); 2 Aug
1957; Sta. 25. Moorea: BPBM 6166; (26, 35–69); 1.5–2.5m.;15 Sep. 1967. CAS 83807; GVF 1308; (13, 42–
63); 30 Apr 1957; Sta. 232. Raiatea: ANSP 85881; (4, 22–53). CAS 83897; GVF 1359; (2, 48–58); 3 Aug
1957; Sta. 27. Tahiti: BPBM 6101; (1, 66); Papara; color, black and white; 22 m; 4 Sep 1967. CAS 83806;
GVF 1339; (32, 23–58); 1July 1957; Sta. 7. CAS 83898; GVF 1344; (4, 51–60); 8 July 1957; Sta. 12. CAS
83801; GVF 1342; (50, 28–63); 4 July 1957; Sta. 10. CAS 83884; GVF 1346; (1, 67); 9 July 1957; Sta. 14.
CAS 83887; GVF 1340; (1, 43); 2 July 1957; Sta. 8. CAS 83802; GVF 1347; (42, 37–61); 10 July; Sta. 15. SU
24586; (2, 28–59); 18 Feb 1929. Tuamotu Archipelago: Raroia: BPBM 9313; (1, 29); Cruise 54, Charles E.
Gilbert; 1 Nov 1961. CAS 83812; GVF 99; (1, 63); 4–8 Aug 1952; Sta. 45. CAS 83962; GVF 103; (3, 62–72);
8 Aug 1952; Sta. 49. Mangareva: USNM 65426; (17, 68–80); 1 Feb 1904-05; Albatross; BPBM 14283; (6,
65–76); 11 Dec 1970. Line Islands: BPBM 7791; (13, 34–48); Fanning I.; 29 Oct 1968. BPBM 14254; (3,
28–46); Kiritimati; 0–2m.; 24 Mar. 1971. CAS 83967; GVF 47; (1, 65); Palmyra I; 18 Aug 1951; Sta. 47.
CAS 83971; GVF 46; (3, 48–68); Palmyra I; 17 Aug 1951; Sta. 46. Rapa: BPBM 12844; (1, 78); 28 Jan 1971.
BPBM 17318; (2, 20–53); 14 Feb 1971.
Diagnosis. A Nectamia lacking dark saddles on body, caudal peduncle with a dark, wide bar, mostly
above the lateral line, caudal fin with dark margins, cheek mark, wide and triangular(Tab. 2); 13 pectoral-fin
rays; total gill rakers usually 26–30 (Tab. 8); body depth 39–44%, caudal-peduncle depth 16–20%, second
anal-spine length 17–21%, and pectoral-fin length 26–29% of standard length (Tab. 3).
Description. Range of proportions (as percentages of standard length) in Table 3 with paratypes and non-
type material in parentheses.
Dorsal fin VII(I)–I,9; anal fin II,8; pectoral fin usually 13-13 rarely 13-14; pelvic fin I,5; principal caudal
rays 9+ 8; pored lateral-line scales 24; transverse scale rows above lateral line 2; transverse scale rows below
lateral line 6; median predorsal scales 3, rarely 2; circumpeduncular scale rows 12 as 5+2+5; total gill rakers
29 (24–31), usually 27 (25–30) well developed, upper arch 2+5 (0-3+5-8), lower arch 22+0 (18-22+0-1).
Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and
vomer; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10+14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural.
Three supraneurals, two supernumerary spines on first dorsal pterygiophore. Basisphenoid present. Supra-
maxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal mar-
gins. Infraorbital shelf present on third bone.
Scales ctenoid on cheek, subopercle, opercle, nape, isthmus, base of pelvic fin with two large weakly
ctenoid scales, no axillary scale, ctenoid scales on base and behind pectoral fin. Pored lateral-line scales com-
plete, fourth scale with two pores above and one pore below main canal.
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TABLE 8. Geographic variation in gill-raker and rudiments counts for Nectamia savayensis.
*= holotype
N=205 Upper Arch Lower Arch Total
6789x 18 19 20 21 22 23 x 24 25 26 27 28 29 30 31 x
INDIAN OCEAN
Zanzibar 1 1 1
Mozambique 25 7.7 421 18.6 2221 26.3
Aldabra I 1 1 1
Seychelle Is 711 7.6 5112 19.9 2 7 8 1 27.4
St Brandon Shoal 1 3 7.8 1 1 1 1 19.5 1 1 1 1 27.3
Mauritius 1 18 7.9 1 2 11 5 20.1 1 3 10 5 28.0
Maldive Is 45 7.6 342 19.9 2322 27.4
Nicobar Is 2 7.0 2 19.0 2 26.0
Cocos-Keeling Is 2 7.0 2 20.0 2 27.0
Sumatra 1 1 1
Ashmore I. 4 9 6.9 2 6 5 18.7 3 8 2 25.6
PACIFIC OCEAN
Taiwan 311 7.8 1283 19.9 2363 27.8
Viet Nam 6 4 1 7.5 3 8 19.7 2 5 3 1 27.3
Thailand 1 1 1
Sulawesi / Seribu 1 1 6.5 1 1 19.5 1 1 26.0
Ambon I 3 3 1 7.7 1 4 2 19.1 1 2 2 1 1 27.9
Banda I. 8 7.0 323 19.0 323 26.0
Luzon / Negros452 6.8 1352 19.7 11333 26.5
Guam / Saipan 6 31 5 8.0 4 8 28 2 20.6 2 5 7 25 2 1 28.5
Enewetak / Rong. 2 12 3 7.1 5 5 7 20.1 6 4 5 2 27.2
Wake 5 2 7.3 3 4 20.6 2 4 1 27.9
Queensland 3817.8 345 20.2 2145 28.0
Bougainville I. 2 5 7.7 3 3 1 21.6 4 3 29.4
Loyalty Island 1 1 1
Rotuma 53 7.4 26 19.8 233 27.1
Tonga Is13817.7 256 20.3 13441 28.1
Fiji Is 7 3 7.3 7 2 20.2 5 3 1 27.6
Samoa 1* 1* 1*
Line Is 1 3 7.8 2 2 20.5 1 1 2 28.3
Mangareva 9 8 7.5 10 6 1 20.5 7 5 4 1 27.9
SUMMARY
E. Africa / Mascar. 12 38 7.8 6 10 26 8 19.7 3 6 13 21 7 27.5
E. I. O./Phil./Indo. 7 31 10 7.1 7 21 17 4 19.4 1 9 14 14 7 3 1 26.5
China Sea 9 16 1 7.7 1 6 16 3 19.8 3 8 11 3 27.6
Guam / Wake I2233647.6 916391 20.5 81116282 128.1
Coral Sea / Line Is1354227.6 940248 20.4 72323216 28.0
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status of Nectamia & 4 new species
Life colors. See Figure 6F. Head, body and fins without stripes; head with broad dark oblique cheek mark
from eye to angle of preopercle, lachrymal and anterior infraorbital with yellowish mark, iris brownish with
white inner ring; head and body a uniform bronze to brownish background becoming silvery below eye and
on abdomen, sometimes with faint pale bars on side or a saddle under the second dorsal fin; caudal peduncle
with a dark basicaudal mark mostly above pored lateral-line scales, absent below pored lateral-line scales,
partial pale region just anterior to peduncle mark; first dorsal fin with dusky distal membranes between spines
II–IV, second dorsal with dusky membranes between the spine and second ray, anal fin with whitish second
spine and first anal ray, pectoral and pelvic rays pale, edge of caudal fin whitish with blackish interior line,
rest of fin pale.
Preserved color pattern. Adults: triangular subocular cheek mark behind upper jaw; first dorsal fin with
dark membrane between spines 1–4; edges of caudal-fin lobes dark; caudal peduncle with an incomplete dark,
wide bar mostly at or above the lateral line, no dark saddles on body (Pacific material occasionally with a sad-
dle under the second dorsal fin); body without pale bars or with faint pale bars on side; stomach and intestine
black; roof of mouth with scattered tiny melanophores not extending onto gill arches. Juveniles: similar to the
adults.
Distribution. Known from the Indian Ocean (absent in the Red Sea) to the Central Pacific (Fig. 13).
Remarks. Kuiter and Kozawa (1999) provide a number of underwater photographs (p. 33, A–E) identi-
fied as Apogon savayensis. Their Apogon species 18 (p34, A–B) may be this species. Günther (1872) clearly
states that there are “...no bands on the body”. No markings were detected on the body from independent
examinations of E. A. Lachner and T. H. Fraser of the type. Some specimens from the West Pacific have a ten-
dency to have a saddle under the second dorsal fin. Average gill-raker counts vary among geographic regions
(Fig. 13, Tab. 8) by 1–1.5 elements. No obvious preserved color pattern differences were observed. This spe-
cies has been confused with Nectamia luxuria which overlaps in geographic distribution except for the west-
ern Indian Ocean (Fig. 7). Nectamia savayensis has a wider cheek mark and almost always lacks pale bars
(when present, faint and few) in contrast to the color pattern of N. luxuria. Lachner (1951, Plate 19a–c) pro-
vided photographs of Nectamia savayensis, but his description included 9 lots of N. luxuria.
Nectamia savayensis has been collected at the stations with N. fusca (ANSP 174727, 174728), N. luxuria
(MCZ 38496, CAS 83800, USNM 341618, 341622, 341626) and N. viria (USNM 263249, 268256, 341613,
341619, 341627). Five specimens in USNM 213113 have buccal enlargement, one with eggs (65 mm SL) and
4 others (64–73 mm SL) without eggs in the mouth. One 57 mm SL specimen in USNM 213187 and 2 speci-
mens in CAS 35369 have buccal enlargement but no eggs were found. The largest of the 1963 specimens was
92 mm SL.
Nectamia similis new species
Figures 3C, 6A, 14–15, 18 and Tables 2–3, 9
Type material: Holotype; USNM 213123 44.0 mm SL; Indonesia, Banda Sea, Kabaena I. VGS 74-1.
Paratypes; USNM 344948 (17, 32–50); same data as the holotype. USNM 344947; (5, 51–59); Indonesia,
Banda Sea, Poka, Ambon I.; 22–23 Jan. 1973, Rumphius Exp. 1, Sta. AB-1; VGS 73-18. ROM 54320; (9, 16–
50); Negros Oriental, Tanton Strait; RW87-29; 19 May 1987; 6–12m.
Additional material: Indonesia: Banda Sea: USNM 213175; (3, 43–52); Ceram, Tandjung Liang; 10
Jan. 1973; Rumphius Exp. 1, Sta. Li-1; VGS 73-6. Java Sea: USNM 213115; (2, 55–60) SW Karimandjawa I;
29 Mar 1974; VGS 74-28. USNM 213120, 213121; (20, 14–42); NW Pulau Tikus, Pulau Seribu; 5 Apr 1974;
VGS 74-34. Molucca Sea: Sulawesi: USNM 112136 (A. 14934, 14935, 23549); (3, 45–54); Talisei I.; 9 Nov
1909; x-ray. USNM 112192 (A.23897); (1, 47); Pendek I.; 15 Dec. 1909. USNM 112189 (A.23847); (1, 49);
Togian I., Gulf of Tomini; 19 Nov. 1909. Banda Sea: USNM 213243; (4, 23–31); Ambon I.; VGS 73-4; 8 Jan.
1973. USNM 210437; (2, 56–59) Ambon I.; VGS 73-20; 24 Jan. 1973. USNM 261157; (15, 16–57); West
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38 · Zootaxa 1691 © 2008 Magnolia Press
Irian, NW tip of Irian Jaya; 2 July 1979. Malaysia: Sabah: USNM 213174; (38, 26–57) Darvel Bay; 1 Feb.
1965; Te Vega Cr 6, Sta. 213. USNM 205668; (1, 50.3); Darvel Bay; 1 Feb. 1965; Te Vega Cr 6, Sta. 213;
cleared & stained. Philippines: ROM 54319; (2, 43–48); Negros Oriental, Siquijor; RW87-20; 14 May 1987;
8–15m. ROM 54320; 9(16–50); Philippines, Negros Oriental, Tanton Strait; RW87-29; 19 May 1987; 6–12m.
ROM 79170; 2(49–57); Negros Oriental, Siquijor I.; RW 87-8; 9 May 1987. SU 38146 (1, 37); Bungau, Sulu
I.; 17 Sept. 1940; A.W. Herre coll. USNM 112130 (A.23869); (1, 47); Rapu Rapu, E. Luzon; 22 June 1909.
USNM 112175 (A.8873); (1, 35); Port Matalvi, Luzon; 23 Nov. 1908. USNM 112133 (A.23776, 23777,
23778, 23779) (4, 44–51); Port Uson, W. of Pinus I., Masbate I.; 17 Dec. 1908. USNM 112134 (A.23768,
23769); (2, 50–51); Ulugan Bay, Palawan I.; 28 Dec. 1908; x-ray. USNM 213610; (1, 37); Machesi I., Pala-
wan Is.; 5 Apr. 1909. USNM 261133; (15, 18–22); Negros Oriental, 09°23'00"N, 123° 115'30"E. South
China Sea: USNM 171387 (A.23356); (1, 46); NW Palawan; Endeavor Str.; 22 Dec. 1909; Papua New
Guinea: CAS 56571; (2, 27–27); Bagaman I., Louisaide Arch.; JEM 85-18; 4 Apr 1985. CAS 65307; (2, 41–
42); Nagada Harbour; SGP 87-7; 24 Apr 1987. CAS 65306; (1, 33); Nagada Harbour; SGP 87-11; 1 May
1987. USNM 261024; (4, 34–54); Hermit I.; VGS 78-19; 4 Nov. 1978. USNM 262406; (1, 41); Kranket I.;
VGS 78-23; 7 Nov. 1978. USNM 213176; (14, 17–52); Bayin Kranket I., Madang Harbor; 30 1970; BBC
1495. BPBM 32430 (2, 48–57); color; Port Moresby, Laloata I.; 27 Oct. 1987. Palau Is.: BPBM 13480; (1,
52); Rmegethu Island; 4.5m.; 5 Dec. 1971. CAS 82661; GVF 1415; (4, 29–54); Auluptagel I; 9 Oct 1957; Sta.
57-37. CAS 84052; GVF 558; (8, 30–55); North Shore of Koror I; 4 Aug. 1955; Sta. 59. CAS 83783; GVF
546; (1, 63); Iwayama Bay, Auluptagel I.; 28 July 1955; Sta. 47. CAS 84051; GVF 526; (33, 24–56); N. side
of Urukthapel I.; 20 July 1955; Sta. 27. CAS 82656; GVF 1414; (31, 18–56); Between Malakal and Aulupta-
gel; 7 Oct. 1957; Sta. 57-36. CAS 84048; GVF 1407; (1, 52) Arakabesan I.; 2 Oct. 1957; Sta. 57-29. CAS
84047; GVF 754; (3, 18–44); Iwayama Bay; 21 Oct. 1955; Sta. 2-44. CAS 82662; GVF 1853; (1, 56); SE of
Arakabesan I.; 22 Jan. 1959; Sta. 59-25. CAS 84049; GVF 532; (12, 22–49); Koror I.; 23 July 1955; Sta. 33.
CAS 84070; GVF 1442; (2, 53–55); Koror I.; 19 Nov 1957. CAS 82655; GVF 1397; (28, 28–62); Babelthaup
I; 23 June 1958; Sta. 58-1. CAS 82663; GVF 1400; (2, 44–57); Babelthaup I.; 25 Sep 1957. CAS 85956; GVF
947; (1, 48); Kayangel I.; 8 Oct 1956. CAS 85945; GVF 639; (14, 24–58); Iwayama Bay; 28 Aug 1955; Yap
I.: CAS 82647; GVF 1904; (1, 44); 9 Dec. 1959; Sta. 81. CAS 84897; GVF 794; (5, 28–36); 5Jul 1956; 0–3m.
Diagnosis. A Nectamia with two dark saddles on body below the first and second dorsal fin, a narrow pale
bar on the opercle reaching above the top of the preopercle, a few narrow pale bars on the body, caudal pedun-
cle with a complete dark wide bar, connected dorsally and ventrally caudal fin with dark margins, subocular
mark, thin and triangular (Tab. 2, Fig. 14); 13 pectoral-fin rays; total gill rakers usually 25–28 (Tab. 9); body
depth 43–45%, caudal-peduncle depth 18–21%, second anal-spine length 19–21%, and pectoral-fin length
26–30% of standard length (Tab. 3).
FIGURE 14. Nectamia similis, holotype, USNM 213123, 44.0 mm SL, Indonesia, Sulawesi, Kabaena I., Tallabassi Bay,
24 Feb 1974, 4.5–5.6 m, VGS 74-1.
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status of Nectamia & 4 new species
Description. For general body shape see Figure 14. Range of proportions (as percentages of standard
length) in Table 3 with paratypes and non-type material in parentheses.
Dorsal fin VII(I)–I,9; anal fin II,8; pectoral fin usually 13-13; pelvic fin I,5; principal caudal rays 9+8;
pored lateral-line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line
6; median predorsal scales 2; circumpeduncular scale rows 12 as 5+2+5; total gill rakers 26 (24–28), well
developed usually 23 (22–27), upper arch 2+5 (0-2+5-7), lower arch 18+1 (17-20+0-2), second arch with
shorter gill rakers 3+18.
Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and
vomer; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10+14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural.
Three supraneurals, two supernumerary spines on first dorsal pterygiophore. Basisphenoid present. Supra-
maxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal mar-
gins. Infraorbital shelf present on third bone.
TABLE 9. Geographic variation in gill-raker and rudiments counts for Nectamia similis.
* = holotype
Scales ctenoid on cheek, subopercle, opercle, nape, isthmus, base of and behind pectoral fin, and two large
weakly Ctenoid scale on base of pelvic fin. Ctenoid lateral-line scales complete, fourth scale with two pores
above main pore and one pore below (Fig. 4C).
Etymology. from the Latin similis meaning like, and refers to the pale bars on the body similar to
Nectamia luxuria and the saddles and caudal mark similar to N. bandanensis. Treated as an adjective.
Life colors. See Figure 6A. Head, body and fins without stripes; head with dark triangular cheek mark
from eye to angle of preopercle, lachrymal and anterior infraorbital with yellowish mark, iris brownish with
white inner ring; head and body bronze to brownish background with broad alternating pale and brown bars;
caudal peduncle with a dark basicaudal mark wider above pored lateral-line scales than below, the bar less
intense below pored lateral-line scales, pale region just anterior to peduncle mark reaching to posterior base of
second dorsal fin and anal fin; first dorsal fin with distal whitish membranes between spines II–IV, pectoral
and pelvic fins pale, second dorsal, anal and caudal fins with whitish edges. Fowler and Bean (1930, p. 43)
record the following field notes for these Albatross specimens (now USNM 112133): “23776 to 23779. Port
Uson, west of Pinas Island. December 17, 1908. Length 61 to 73 mm. Iridescent silvery pink. Top of head and
3 saddles across back blackish olive, one saddle below each dorsal and one on hind part of caudal peduncle,
last darkest. Triangular bronze blotch below eye, pointing obliquely across cheek. Opercle bronze, with 2 bro-
ken silvery vertical bars. Similar indistinct bars on side of body, about 10 between head and axil of dorsal.
Chin and lower surface of head paler than above. Fins similar to adjacent body but paler. Caudal with 2 dusky
regions on each lobe one basal and one median.”
N=79 Lower Arch Upper Arch Total
67 8x 18 19 20 x 24 25 26 27 28 x
Palau / Yap 1 8 3 7.2 1 6 5 19.3 2 3 6 1 26.5
Philippine 5 7.0 3 2 18.4 3 2 25.4
Indonesia 2 24* 6.9 5* 18 3 18.9 1 5 17* 3 25.8
Irian Jaya 2 11 1 6.9 6 6 2 18.7 1 6 4 3 25.6
Papua New Guinea 4 17 1 6.9 2 12 8 19.3 1 4 9 7 1 26.1
Total 9 65 5 6.9 17 44 18 19.0 3 20 35 19 2 26.0
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FIGURE 15. Distribution of material examined for Nectamia similis .
Preserved color pattern. Holotype: Head with faint dark mark on lachrymal and anterior infraorbitals,
well-defined dark, angular cheek mark ending in a point at edge of preopercular ridge, short pale bars, one
behind and along cheek mark, a second above preopercle arm, a third on opercle, no other markings on head;
body with alternating wider darker bars and narrower well-defined lighter bars from opercular flap to near
posterior origin of second dorsal fin, caudal peduncle with a wide lighter bar followed by a complete darkish
bar, more intense above pored lateral-line scales than below, darkish dorsal saddles under first and second dor-
sal fins, sometime variable under first dorsal fin; first dorsal fin dusky, second dorsal fin, anal fin and pelvic
fins dusky to pale, pectoral fins pale, caudal fin darkish on several of outer rays forming dark edges, interior of
caudal fin pale (Fig. 14); roof of mouth and upper gill arches with numerous scattered melanophores; stomach
and intestine blackish. Adults: Similar to holotype with saddle under first dorsal usually present, dark caudal-
peduncle bar more intense above pored lateral-line scales Juveniles: similar to the adults except that the dark
caudal bar with uniform intensity throughout.
Distribution. Known only from the West Pacific to Yap (Fig 15).
Remarks. Nectamia similis appears somewhat similar to N. luxuria based on the pale bars and dark-edged
caudal, but differs by having dorsal saddles, a more complete caudal-peduncle bar, pale bars on the opercle, a
narrower cheek mark, fewer pale body bars and slightly lower gill-raker counts. The dorsal saddles and lower
gill-raker counts are more like N. bandanensis. The cleared and stained specimen reported as Apogon bandan-
ensis by Fraser (1972) is Nectamia similis.
This species has been taken with Nectamia luxuria (USMN 112135, 112161, 112170, 169723, 171400,
209644, 341634, 344924) and N. viria (USNM 210492, 213184, 213341, 341633). The largest of 323 speci-
mens was 63 mm SL.
Zootaxa 1691 © 2008 Magnolia Press · 41
status of Nectamia & 4 new species
Nectamia viria new species
Figures 3B, 6C, 7, 16, 18 and Tables 2–3, 10
Type material: Holotype Apogon viria USNM 319149 57.4 mm SL, Philippine Is., Sangay Siapo Is., 5–15
feet, 20 May 1988, Sta. EM 88-9, E. Menez and party, x-ray. Paratypes; USMN 344935 (136, 18–58 mm SL),
same data as the holotype. Paratypes ROM 54324; (14, 41–53); Cebu, Similon I., Rohol Straits; RW87-37; 21
May 1987; 4–8m.
Additional material: West Pacific Ocean: Indonesia: Banda Sea: USNM 263249; (22, 52–72); Banda
Is., Naira I.; VGS 74-9; 8 Mar. 1974. USNM 213111; (9, 16–47); Banda Is., Naira I.; VGS 74-4; 7 Mar. 1974.
USNM 210960; (23, 28–60); Nusa Laut; 16 Jan 1973; Rumphius Exp. I, Sta. NU-1. USNM 339756; (1, 42)
and USNM 341624; (23, 37–72); Ambon; 14 Mar 1974; VGS 74-13. USNM 261140; (4, 15–24). Kai Ids.,
5°31'36"S, 132°17'48"E; USNM 210492; (5, 44–57); Poka, Ambon I.; 22–23 Jan. 1973, Rumphius Exp. 1,
Sta. AB-1; VGS 73-18. Java Sea: USNM 213116; (348, 10–49); Pulau Seribu; 4 Apr 1974; VGS 74-31.
USNM 213117; (15, 15–21); Pulau Seribu; 4 Apr 1974; VGS 74-32. USNM 209695; (2, 26–29) and USNM
213184; (3, 26–29); Ceram, Tandjung Liang; 10 Jan 1973; Rumphius Exp 1, Sta Li-1; VGS 73-6. Malaysia:
Sabah: USNM 213341; (104, 25–53); Darvel Bay; 2 Feb. 1965; Te Vega Cr 6 Sta. 216. Philippines: ANSP
174730; (18, 28–51); CMP-P; Nov 1917. ANSP 77366; (2, 31–51); CMP-P; Nov 1917. CAS 49608; (32, 36–
56); Layia, Luzon I.; 28 Jun 1948. SU 69752; (13, 18-32); Sitankay, Sibutu Is.; 9 Aug 1931. ROM 79171;
1(51); Negros Oriental, Siquijor; RW87-20; 14 May 1987; 8–15m. ROM 79169; 1(48); Negros Oriental,
Siquijor I.; RW 87-20; 9 May 1987. SU 69753; (5, 43–50); Sitankay, Sibutu Is.; A.W. Herre; 7 Jan 1937.
USNM 268271; (4, 36–62); Marinduque I.; 24 Apr. 1908. USNM 153237 (A.23559, 23560); (9, 46–53);
Romblon I.; 26 Mar 1908. USNM 171380; (3, 26–49); Palawan I.; 11 Apr. 1909. USMN 341619; (7, 32–52);
Palawan Is., Bararin I., SP 78-20; 23 May 1978. USNM 261137; (27, 8–21); Pan de Azucar I., 10°57'48"N,
121°13'32"E. USNM 261135; (8, 9–24); Cuyo I. USNM 341627; (8, 39–54); Pamilikan I., 9°29' 20"N,
123°55'00"E; SP 78-43; 12 June 1978. USNM 261139; (37, 9–21); eastern Negros I. 9°4'00"N, 123°8'48"E.
USNM 268256; (ca. 250, 17–68); Negros Oriental, Siquijor I.; LK 79-15, 15 May 1979. USNM 268260; (3,
50–62); Mindanao, Zamboanga del Norte; LK 79-9; 4 May 1979. USNM 329214; (9, 16–53); Mindanao,
Zamboanga del Norte; LK 79-10; 4 May 1979. USNM 268257;(ca. 100, 15–70); Mindanao, Zamboanga del
Norte; LK 79-7; 3 May 1979. USNM 261149; (8, 50–61); Negros I., south tip; SP 78-11; 13 May 1978.
USNM 314515; (85, 12–51); Sibutu I.; EM 88-6; 26 May 1988. USNM 319152; (16, 15–44); Sibutu I.; EM
88-7; 27 May 1988. SU 38147; (10, 18–60); Jolo I., Sulu Is.; 16 Sep 1940. USNM 112177 (A.8695); (1, 48);
Jolo I.; 19 Sep. 1909. USNM 171376; (2, 37–46); Tonquil I.; 14 Sep. 1909. Papua New Guinea: USNM
341633; (8, 14–42); Hermit I.; VGS 78-19; 4 Nov. 1978. USNM 341613; (3, 51–53); Hermit Is., Amot I.;
VGS 78-10; 30 Oct. 1978. USNM 261156; (10, 38–53); Louisade Arch., Panapompom I.; BBC-1693, 15 June
1979. USNM 213181; (7, 37–57); Madang Harbor; 26 May 1970; BBC-1488 & 1490. USNM 213182; (6, 43–
50); Madang, S. end Belian I; 22 May 1970; BBC 1478. USNM 268271; (4, 36–50); Kranket I.; VGS 78-22;
7 Nov. 1978. USNM 296841; (1, 42); Nagada Harbour; SGP 87-17; 8 Apr. 1987. Bismark Archipelago:
USNM 213183; (6, 32–41); Rabaul, New Britain; 28 Feb 1965; Te Vega Cr. 6, Sta. 236. USNM 268264; (29,
27–39); Keraward I, Duke of Kirk Gp.; 25 Feb 1965; Te Vega Cr. 6, Sta. 234. New Caledonia: ROM 65765;
(3, 59–63); Recif Mbere, 22°20'30"S 166°14'45"E; RW 91-19; 11 Sep 1991; 3–6 m. ROM 65772; (1, 59);
Recif Laregnere, 22°20'00"S 166°17'50"E; RW 91-13; 6 Sep 1991; 0–9 m. ROM 65798; (2, 56–67); Recif
Laregnere, 22°20'00"S 166°17'50"E; RW 91-12; 6 Sep 1991; 8–11 m. ROM 65810; (16, 44–67); Isle Nge,
22°19' 40"S 166°18'20"E; RW 91-11; 5 Sep 1991; 5–6 m. ROM 65812; (1, 62); Isle Nge, 22°19' 40"S
166°18'20"E; RW 91-11; 5 Sep 1991; 5–6 m. ROM 65929; (1, 57); Isle Te Ndu, 22°18'30"S 166°17'00"E; RW
91-3; 30 Aug 1991; 3–6 m. ROM 65931; (4, 36–55); Ua, 22°42'30"S 166°48'20"E; RW 91-22; 12 Sep 1991;
0–11 m. ROM 65932; (1, 61); Isle de Crouy, 22°21'00"S 166°21'00"E; RW 91-29; 16 Sep 1991; 3–6 m.
USNM 324821; (1, 58); Noumea, 22°48'30"S 166°27'24"N; 7 Nov 1991; JTW 91-1. Australia: Queensland:
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ANSP 123338; (27, 21–54); Little Hope Island; TSA-18; 17 Jan 1969. ANSP 123400; (2, 51–57); Endeavour
Reef; TSA-4; 5 Jan 1969. ANSP 174729; (17, 21–30); Endeavour Reef; TSA-17; 16 Jan 1969. ROM 39806;
(1, 50); Eagle I.; RW 81-11; 19 Sep 1981. ROM 39820; (2, 54–59); Yonge Reef; RW 81-10, 18 Sep 1981.
ROM 44266; (2, 55–64); Escape Reef; ER 81-32; 6 Nov 1981. One Tree I.: USNM 213126; (4, 53–69),
USNM 213127; (8, 51–60); 25 Nov. 1966; VGS 66-8. USNM 213128 (1, 49), 213129; (7, 44–65); VGS 66-
14; 1 Dec. 1966. WAM P24725; (3, 41–58); Lizard I; 9 May 1974. Palau Islands: CAS 84050; GVF 529; (3,
30–42); Iwayama Bay; 22 July 1955; Sta. 30. CAS 98026; GVF 1414; (5, 52–53); Between Malakal and
Auluptagel; 7 Oct. 1957; Sta. 57-36. CAS 82657; GVF 536; (7, 31–53); between Peleliu & Ngedebus; 25 July
1955; Sta. 37. CAS 84062; GVF 1829; (5, 28–44); Babelthaup I; 23 June 1958; Sta. 58-1. CAS 98027; GVF
1415; (4, 46–50); Auluptagel I.; 9 Oct 1957; Sta. 57-37 CAS 98025; GVF 754; (2, 25–32); Iwayama Bay,
Auluptagel I. SU 29157; (5, 19–59); 12–18 Oct 1933. CAS 95834; (1, 38); Malakal I.; 19 May 1997; photo-
graph. Yap I.: CAS 84066; GVF 1910; (31–58); 16 Dec 1959; Sta. 87. CAS 84095; GVF 1935; (5, 34–52); 6
Jan 1960. Ifaluk Atoll: CAS 82653; GVF 221; (2, 57–70); 16 May 1960. Pohnpei: CAS 98024; GVF 493; (3,
37–39); 4 Oct 1954. Solomon Islands: ROM 42942; (2, 18–29); Guadalcanal, Honiara; 4 May 1987; 21 m.
SU 25143; (3, 46–55); Ugi I.;10 Apr 1929. USNM 358404; (1, 22); Santa Cruz Is, Lomlom I; 15 Sep 1998;
SOL 98-3. Vanuatu:12 May 1997; Espiritu Santo Island, W. of Luganville, 15°31'30"S 167°09'32"E; 0–5 m.
Fiji.: ROM 43264; (1, 35); Great Astrolabe Reef; WE 83-37; 2 Apr 1983.
TABLE 10. Geographic variation in gill-raker and rudiments counts for Nectamia viria.
*= holotype
Diagnosis. A Nectamia with one dark saddle on body below the second dorsal fin, caudal peduncle with a
complete dark wide bar, connected dorsally and ventrally caudal fin without dark margins, subocular mark,
thin and triangular (Tab. 2, Fig. 13), 13 pectoral-fin rays; total gill rakers usually 25–28 (Tab. 10); body depth
N=170 Upper Arch Lower Arch Total
678x 18 19 20 21 22 x 24 25 26 27 28 29 x
Ifalik / Pohnpei 3 2 7.4 1 4 19.8 1 2 2 27.2
Palau / Yap 4 17 6.8 2 7 10 2 20.6 1 2 7 10 1 27.4
Philippines 12* 7 6.4 3 11 4* 1 19.2 2 7 8* 2 25.5
Sabah, Darvel Bay 8 7 6.5 1 8 6 19.3 5 8 2 25.8
Indonesia, Seribu I. 13 7 6.1 1 9 5 19.3 1 7 7 25.4
Indo., Ambon / Nusa Laut 6 9 6.6 7 8 19.5 4 5 6 26.1
Indonesia, Banda I. 9 6 7.4 1 7 7 20.4 1 5 5 4 27.8
New Guinea, Keraward I. 1 8 6.9 2 4 3 20.1 1 1 4 3 27.0
New Guinea, Madang 5 1 7.2 3 3 20.5 2 4 27.7
Australia 1 7167.6 2157 20.2 1 6134 27.8
New Caledonia 5 9 7.6 1 9 4 20.2 1 3 7 3 27.9
Solomon I. 1 1 1
Fiji 1 1 1
Vanuatu 1 8 1 7.0 7 3 20.3 1 6 2 1 27.3
SUMMARY
Palau / Pohnpei 4 20 2 6.9 3 11 10 2 20.4 1 3 9 12 1 27.3
Philippines- Ambon - Seribu 39 25 6.4 5 35 23 1 19.3 3 23 28 10 25.7
Banda I. - Fiji 3 44 33 7.4 6 47 27 20.3 1 5 28 34 12 27.6
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status of Nectamia & 4 new species
44–47%, caudal-peduncle depth 17–19%, second anal-spine length 16–18%, and pectoral-fin length 26–29%
of standard length (Tab 3).
Description. For general body shape see Figure 16. Range of proportions (as percentage of standard
length) in Table 3 with paratypes and non-type material in parentheses.
Dorsal fin VII(I)–I,9; anal fin II,8; pectoral-fin 13-13; pelvic fin I,5; principal caudal rays 9+8; pored lat-
eral-line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line 6;
median predorsal scales 2; circumpeduncular scale rows 12 as 5+2+5; total gill rakers 26 (24–29), well devel-
oped 25 (23–29), upper arch 1+5 (0-2+5-6), lower arch 20+0 (18-20+0-1), second arch with shorter gill rakers
3+18.
Villiform teeth in several rows on the premaxilla; two rows on the dentary; one row on the palatine and
vomer; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10+14. Five free hypurals, one pair of slender uroneurals, three epurals, a free parhypural.
Three supraneurals, two supernumerary spines on first dorsal pterygiophore. Basisphenoid present. Supra-
maxilla absent. Posttemporal serrate on posterior margin. Preopercle serrate on vertical and horizontal mar-
gins. Infraorbital shelf present on third bone.
Scales ctenoid on cheek, subopercle, opercle (except posterior scale cycloid), nape, isthmus, base and
behind pectoral fin, base of pelvic fin with two large ctenoid scales. No axillary scale, large scale with ventral
edge bent. Pored lateral-line scales complete, fourth scale with one pore above and one pore below main pore
(Fig. 4B).
FIGURE 16. Apogon viria, holotype, USNM 319149, 57.4 mm SL, Philippine Is., Sangay Siapo I., 28 May 1988, 1.5–
4.5 m, Philippine Expedition, EM 88-9.
Etymology. from the Latin viria meaning bracelet and refers to the band around the caudal peduncle.
Treated as a noun in apposition.
Life colors. See Figure 6C. Head, body and fins without stripes; head with dark narrow triangular cheek
mark from eye to angle of preopercle, lachrymal and anterior infraorbital with yellowish mark, iris brownish
with white inner ring; head and body bronze to brownish background occasionally with a few faint pale bars,
saddle under second dorsal fin; caudal peduncle with a complete dark caudal-peduncle mark, the bar less
intense below pored lateral-line scales, pale region just anterior to peduncle mark not reaching to posterior
base of second dorsal fin but reaching anal fin; first and second dorsal fins uniform dusky, pectoral pelvic and
caudal fins pale. Kuiter and Kozawa (1999) provide four of underwater photographs of this species (p. 33, A–
C, E) identified as Apogon bandanensis. .
Preserved color pattern. Holotype: Head with faint dark mark on lachrymal and anterior infraorbitals,
well-defined dark, angular cheek mark ending in a broad point at edge of preopercular ridge, no other mark-
ings on head; body without alternating darker bars and well-defined lighter bars from opercular flap to near
posterior origin of second dorsal fin, caudal peduncle with a wide lighter bar followed by a complete darkish
bar of nearly uniform intensity, darkish dorsal saddle under second dorsal, saddle not apparent under first dor-
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sal fin; first dorsal fin dusky, second dorsal fin, anal fin and pelvic fins dusky to pale, pectoral fins pale, caudal
fin uniform darkish (Fig. 16); roof of mouth and upper gill arches pale, without numerous scattered melano-
phores; stomach and intestine blackish. Juveniles: similar to the adults.
Distribution. Known from Indonesia, Philippines, Australia and out to Fiji (Fig. 7).
Remarks. Specimens from the Fiji, the Coral Sea and Banda Island have higher gill-raker counts than
those from the Philippines, Sabah and Seribu Island (Tab. 10, Fig 7). Material from Ambon, Nusa Laut and
Pulau Banda, all in the Banda Sea, about 158 km apart had specimens with differing counts (Fig. 7, Tab. 10).
No other differences were noted. This species has a similar dark caudal band and a bright white band just ante-
riorly similar to the caudal-peduncle pattern of Nectamia annularis. These two species differ in the shape of
the cheek mark, narrower in Nectamia viria, and the presence of a saddle below the second dorsal fin in N.
viria.
This species has been taken at stations with one or more of the following species: Nectamia fusca (USMN
341625, ANSP 123440), N. luxuria (ANSP 77377, USNM 112132, 169727, 213158, 213159, 341617,
341620, 341621, 341626, 341629), N. savayensis (USNM 213113, 261151, 262189, 329215, 341627) and N.
similis (CAS 62656, USNM 261024, 344947). One 65mm SL specimen contained eggs in the mouth and four
other specimens has buccal enlargement (USNM 263249). The largest of 1607 specimens was 72 mm SL.
Nectamia zebrinus (Fraser, Randall and Lachner, 1999)
Figures 6D, 13, 17, 18 and Table 2–3, 11
FIGURE 17. Nectamia zebrinus, holotype, USNM 213422, 59.6 mm SL, N. Of Ras Burqa, N.W. coast of Aqaba, Egypt,
23 July 1969, to 10 m., VGS 69-7.
Type material: Holotype Apogon zebrinus USNM 213422; 59.6 mm SL; N. Of Ras Burqa, N.W. coast of
Aqaba, Egypt; 23 July 1969; to 10 m.; Victor G. Springer; VGS 69-7; x-rayed. Paratypes: Red Sea: Egypt:
BPBM 31807 was USNM 213424; (4, 57–71); Gulf of Aqaba, Ras Muhammad; 0 to 10m.; 26 Sep. 1969;
VGS 69-28. USNM 341635; (4, 49–73); data same as holotype. TAU P.9672; (1, 67); Gulf of Aqaba; NS
4181; 8 Oct. 1968. Sudan: BPBM 19741; (2, 38–44); Suakin Harbor, 11 Oct. 1974. BPBM 27417; (1, 41);
color photo; just N. of Port Sudan, 9 Jan. 1980. Eritrea: USNM 213423 (1, 47); N. End of Isola Delemme, off
Massawa; 7 Aug. 1969; VGS 69-9. Saudi Arabia: BPBM 30388; (4, 58–74); 29 km. S. of Yanbu, 30 May
1984. Yemen: USNM 213421 (15, 55–63); Gold Mohur Bay; 21 Dec. 1964; IIOE Cr. 9; F. Talbot Sta. 26.
Additional material: Red Sea: Israel: TAU P.9671; (1, 27); Gulf of Aqaba. HUJ 12007; (3, 17–26); Elat;
HUJ 11984; (2, 16–25); Elat; HUJ 12859; (1, 20); Dahab.
This species is not treated in detail, see Fraser et al. 1999 for more information.
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status of Nectamia & 4 new species
Diagnosis. A Nectamia with pale vertical bars on body, basicaudal mark below lateral-line scales in
adults, caudal fin with dark margins, subocular mark broad, triangular (Tab. 2, Fig. 17); 13 pectoral-fin rays;
total gill rakers usually 25–28 (Tab. 11); body depth 40–47%, caudal-peduncle depth 18–22%, second anal-
spine length 19–23%, and pectoral-fin length 26–32% of standard length (Tab. 3).
Additional description. Holotype with ctenoid scales on cheek, nape and opercle, cycloid on side of isth-
mus, ctenoid ventrally, cycloid on base of pectoral fin, ctenoid behind pectoral fin, ctenoid on rest of body,
fourth lateral-line scale with 2 pores above main pore and 2 pores below, base of pelvic fin missing pelvic
scales, large scale present at side of pelvic spine with a narrow bend along ventral edge, not produced.
Paratypes with two large ctenoid scales on base of pelvic fin.
Second arch with developed gill rakers, 3+ 16 and 2 rudiments on lower limb.
TABLE 11. Gill-raker and rudiments counts for Nectamia zebrinus.
* = holotype
Life color. See Figure 6D. Head, body and fins without stripes; head with broad dark triangular cheek
mark from eye to angle of preopercle, lachrymal and anterior infraorbital with yellowish mark, iris brownish
with white inner ring; head and body bronze to brownish background with pale bars; caudal peduncle with a
dark caudal-peduncle mark wider above pored lateral-line scales than below, the bar less intense below pored
lateral-line scales, pale region just anterior to peduncle mark not reaching to posterior base of second dorsal
and anal fins; first dorsal fin with distal darkish membranes between spines II–IV, second dorsal fin with dark-
ish anterior edge, anal fin with whitish anterior edge, pectoral and pelvic fins pale, and caudal fin with faint
whitish edges. Kuiter and Kozawa (1999) provide a three underwater photographs of this species (p. 34, A–C)
identified as Apogon species 21.
Preserved color pattern. Adults: wide, triangular subocular cheek mark behind upper jaw; first dorsal fin
with dark membrane between spines 1–4; edges of caudal lobes dark; caudal peduncle with a nearly complete
dark wide bar, connected dorsally and almost connected ventrally; pale, slightly wavy bars present on body
from below the first dorsal fin and behind the pectoral fin to just behind the ends of the second dorsal fin base
and anal fin base, bars extend above the lateral-line scales starting about the second dorsal fin; no dark saddle
under the first or second dorsal fin (Fig. 17); stomach and intestine black. Juveniles: similar to the adults,
except with a complete, dark caudal-peduncle band.
Distribution. Known only from the Red Sea and the Gulf of Aden (Fig. 13).
Remarks. Rather bold pale bars on the body and a dark caudal-peduncle mark is not quite complete along
the anterio-ventral part of the band will identify Nectamia zebrinus from N. annularis and N. savayensis (Tab.
2). Caudal-peduncle depth is slightly greater in Nectamia zebrinus than in N. annularis. Indian Ocean
Nectamia savayensis may be distinguished from N. zebrinus by the lack of well developed bars on the side, an
incomplete caudal mark almost always at or above the lateral line (see Winterbottom et al. 1989, fig. 164)
even in juveniles and a shorter second anal spine (14–20% of SL). The gill-raker counts show, at best, slight
modal or average differences, particularly the lower gill arch (Tabs. 8 and 11) between these two species.
Nectamia luxuria has more pale bars, yellow edges to the vertical fins and higher number of gill rakers.
This species has been collected at the same stations with Nectamia annularis (BPBM 20373, 30386,
USMN 212431)) in the Red Sea and Gulf of Aden (USMN 212447) and with N. fusca (USNM 212882). The
largest of 40 specimens was 74 mm SL.
N=37 Upper Arch Lower Arch Total
67 8 x 17 18 19 20 x 24 25 26 27 28 x
Red Sea 1 21* 15 7.4 1 7 16 13* 19.1 1 5 14 9* 8 26.5
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TABLE 12. Coding of character for the phylogenetic analysis using PAUP. Holapogon maximus was used as the out-
group.
Color as a basis for species relationships
Color patterns within groups of cardinalfishes show variations of specific patterns. Determining a template to
polarize pattern trends is clearly associated with some assumptions. Markings and patterns of color commonly
appear across apogonids, for example, caudal-peduncle markings, body stripes, bars, spots and ocellae; head
stripes, bars, spots and ocellae; fin stripes, spots and ocellae. Some color patterns change with ontogeny, day
versus night and breeding behavior. Such issues dampen the idea of using color patterns as containing useful
phylogenetic information: however, Mabuchi et al. (2006) found a strong molecular basis for color patterns
Character State Code Character State Code
caudal fin roof of
marking uniform with edge 0 mouth pale 0
darkish edged 1 speckled melanophores 1
yellowish edged 2 dark 2
whitish edged 3 upper
cheek gill arches pale 0
mark broad through eye dorsally 0 with melanophores 1
wide triangle 1 juvenile
narrow triangle 2 to adult spot spot 0
narrow line 3 peduncle uniform uniform 1
body bar mark(s) spot ½ band 2
marks none 0 ½ band ½ band 3
few faint bar(s) or dark bar(s) 1 complete band >3/4 band 4
pale bars well apart 2 band band 5
narrower dark/pale bars 3 stomach
pale lines within dark bars 4 and gut both pale 0
body both black 1
saddles none or faint 0 pectoral
1 dark saddle 1 fin-rays 14 0
2 dark saddles 2 13 1
lachrymal lateral
mark uniform 0 line tree-like above & below 0
yellowish mark 1 scale > 3 main pores above 1
pores 2 main pores above 2
1 main pore above 3
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status of Nectamia & 4 new species
and phylogeny within Apogon, sensu lato, providing evidence that some basic patterns may have evolved
more that 20 million years before present. They used two species of Fowleria as the outgroup. Holapogon
maximus was used as the outgroup for color patterns and morphologic characters. This species is hypothesized
to have retained more osteological characteristics of the common ancestor shared with other apogonids and
has fewer specializations than Coranthus or Glossamia. The color pattern of Holapogon (Fraser, 1973, fig. 1;
Kyushin et al. 1977, fig. 115; Randall, 1995, fig. 405; Kuiter and Kozawa, 1999, p.91) has dark spots all over
the body and two bars on the head, one through the eye forming a cheek mark and the other extending from
the nape onto the opercle. The vertical fins and pelvic fins were mostly uniform with the membranes between
the first and third dorsal spines darkish. The fifth ray of the pelvic fin had alternating white and darkish areas.
At least three other species in Apogon, sensu lato, have spots over the body (Apogon maculiferus, a Hawaiian
endemic, A. regani a Mascarene Plateau endemic, and Apogon euspilotus, known only from New Caledonia).
Such overall spotted color patterns could lead to all of the known pattern variations in cardinalfishes.
Adult color patterns, except for the caudal-peduncle mark where the change from juvenile to adult, were
used in this analysis. Elaborations of different patterns and color shared by species should provide evidence of
kinship. A brownish/bronze coloration of the head and body within Nectamia was chosen as the basal color
pattern for pattern trends, much like Nectamia fusca.
TABLE 13. Character matrix used for the phylogenetic analysis using PAUP for Nectamia. The outgroup was Hola-
pogon maximus. Apogon melas (group II after Mabuchi et al. 2006) was treated as an ingroup with the species of
Nectamia.
The heads of all species lacked color patterns other than a dark cheek mark on all species, a lachrymal and
infraorbitals with a yellowish dash (known for eight of nine species), and a pale opercular bar in one of nine
species. The shape of the single cheek mark varied among the species from a thin line, a narrow triangle-like
dash or a broad dash, often triangle-like. A caudal peduncular mark was present on all species. The shape,
extent and intensity of the mark varied among the species. The shape ranged from an off-center spot not con-
necting dorsally, an intense off-center bar connected dorsally, a narrower, less intense partial bar, or a com-
plete bar of similar intensity around the caudal peduncle. Dorsal saddles may be absent under the first and
second dorsal fins, to present under the second dorsal or present under both dorsal fins. Pale body bars may be
caudal pale roof upper yg-ad stomach
Characters fin cheek body dorsal lachrymal of gill peduncle and gut pectoral pored
edge mark bars saddles mark mouth arch marks color fin rays scales
123 4 5 6 7 8 9 10 11
Species
H. maximus 00000000000
A. melas 03000001003
N. ignitops 02121215112
N. annularis 31101005113
N. viria 02111005113
N. similis 32221114113
N. fusca 03101000113
N. savayensis 11101003112
N. zebrinus 11301004113
N. luxuria 21411003113
N. bandanensis 02121215113
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absent, faintly present, always present, few or many in number. A pale bar may be present or absent on the
opercle. The iris of the eye may be darkish or reddish. Melanophores making the roof of the mouth dark and
gill arches occur in some species. Stomach and intestine were blackish with pale peritoneum in all species.
Morphologic characteristics were examined to provide support for hypothesized relationships. There was little
or no variation in meristics or morphometry among species of Nectamia, except for body depth and gill-raker
counts. Mabuchi et al. (2006) recovered Apogon melas as a sister species to two species of Nectamia and
placed Ostorhinchus aureus, a close relative of O. fleurieu, in a different lineage. Selected characteristics were
coded (Tabs. 12 and 13) for all species used in PAUP for hypothesized relationships within Nectamia (Fig.
18).
Discussion
Species of Nectamia have 13 pectoral-fin rays, eight anal rays, high first arch total gill-raker counts (24–31),
developed gill rakers on second arch, black stomach and intestine, three supraneurals, two supernumerary
spines on the first dorsal pterygiophore, eight visible dorsal spines, a pair of slender uroneurals, three epurals,
five free hypurals, forked caudal fin, all scales ctenoid and a relatively simple pored scale architecture on the
lateral-line scales. Color pattern and shape combinations relating to cheek marks, pale barring on body, sad-
dles under each dorsal fin, caudal-peduncle marking and edging of the caudal fin margins best distinguish the
species complex (Table 1). Juvenile color patterns differs mostly in the development of the caudal-peduncle
marking.
Apogon melas shares cheek mark and scale pore characteristics with Nectamia but the other characteris-
tics differ: body uniformly dark in adults, 14 pectoral rays, low number of gill rakers, ocellus or dark marks in
the second dorsal and anal fins, pale stomach and intestine, as well as, an unusual dark/light barring juvenile
color pattern. Cheek marks and/or dorsal saddles are found in other Apogon, sensu lato, such as, Apogon timo-
rensis and Apogon gardineri but these species have much lower gill-raker counts and more than 13 pectoral
fin-rays. The above discussion involved species Mabuchi et al. (2006) treated in their ‘Ostorhinchus II’ or
likely to be placed there in my opinion.
At least the species that have been associated with the type of Ostorhinchus (Gon, 1987; Randall et al.
1990) have color pattern and morphological characters that differ from Nectamia: fused sixth radial, loss of
eighth dorsal spine, fewer developed gill rakers on the second gill arch, a prominent stripe on the snout
through the eye, a maxilla stripe, no bars or saddles on the body. Other cardinalfishes in the subgenus
Ostorhinchus with higher gill-raker counts include species in the Apogon notatus group, however these spe-
cies have 14 or 15 pectoral-fin rays, a different head shape, dark stripe(s) present or absent on the body, head
stripes or linear disconnected markings (snout stripe), dark spots on predorsal/nape area and/or body present
or absent and a well-defined midlateral basicaudal spot present. Mabuchi et al. (2006) recovered a clade they
identified as ‘Ostorhinchus group III’ with head and/or body stripes that included the notatus group.
The cladistic tree reported here suggests that the molecular basis for the use of color as a phyletic tool
used at higher taxa levels by Mabuchi et al. (2006) can be used for hypotheses of relationships among species
of a given clade. Meristics, morphometrics and osteology have little variation among the species of Nectamia
which might provide phyletic inferences at the species level. Nine of the eleven characters were based on
color patterns (Tables 12 and 13).
Randall (1998) recently reviewed shore-fish zoogeography of the Indo-Pacific region. The known distri-
bution patterns and other discussed issues, such as geminate species, have relevance given the geographic
variation found in species of Nectamia. Species of Nectamia are represented by one or more species in almost
all Indo-Pacific regions. The Hawaiian chain of islands, Johnston Island, the Marquesas Islands, Easter Island,
Lord Howe, Norfolk and Kermadec Islands in the Pacific Ocean and the Persian Gulf - northern Arabian Sea
Zootaxa 1691 © 2008 Magnolia Press · 49
status of Nectamia & 4 new species
to Somalia in the Indian Ocean do not have known reliable reports or recent collections of this group.
Although there are few reported cardinalfish collections from the Nicobar and Andaman Islands in the Indian
Ocean and from Nauru, Tarawa, Funafuti areas in the Pacific Ocean, representatives of this genus should be
present.
FIGURE 18. A hypothesis for phylogenetic relationships within Nectamia based characteristics in Tables 12 and 13.
One of six best trees retained based on a heuristic search using parsimony was identical to the 50% majority rule consen-
sus of 6664 trees. All 11 characters of equal weight were parsimony-informative. Holapogon maximus was used as an
outgroup.
Nectamia fusca is absent from the Mascarene Plateau even though it has been taken from coastal Africa,
Madagascar, Comores and the Maldives, and has not been reported from relatively well-collected areas of the
Indian Ocean: Chagos Archipelago, Mauritius, Rodrigues, and Réunion. Nectamia savayensis has not been
reported from the Red Sea. Nectamia luxuria reaches the Maldives in the Indian Ocean, but has not been
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50 · Zootaxa 1691 © 2008 Magnolia Press
reported from the western Indian Ocean and may have had its most recent ancestor in common with the Red
Sea species, N. zebrinus (Fig 18). The South China Sea species, Nectamia ignitops may have had its most
recent ancestor in common with N. bandanensis. Nectamia viria, N. bandanensis and N. luxuria show a por-
tion of their distributions consistent with the Carolina Islands conduit (Springer, 1982) on to the Pacific Plate.
Three species, Nectamia fusca, N. luxuria and N. savayensis extend onto the Pacific Plate outward from the
Samoan Islands.
Gill raker variation shows a consistent geographic pattern of lower counts in the Philippines or Philip-
pino-Indonesian area as compared with adjacent areas for Nectamia fusca, N. luxuria, N. savayensis and N.
viria (Tabs. 5, 7–8, 10, Figs. 9, 12, 13 and 16). Nectamia luxuria has a cline of increase number of gill rakers
from the central Philippino-Indonesian area of lower numbers westward to the southern China Sea and highest
numbers in the Indian Ocean to the Maldives (Fig. 12). Another increasing cline in gill raker counts extends
from the Coral Sea eastward to beyond Samoa. Lastly, this species may have another increasing cline extend-
ing out along the Carolina Islands. Nectamia viria has strong gill-raker count differentiation, at the level often
accepted for species, with lower gill-raker counts centered in the Philippine-Seribu region and higher counts
to the east along the Carolina Conduit and to the south in the area of the Banda Is., northern New Guinea,
Coral Sea to Vanuatu and Fiji. Intermediate gill-raker counts for specimens of N. viria were taken at Ambon
and Nusa Laut, and high raker count specimens were taken at Pulau Banda, about 158 kilometers apart in the
Banda Sea (Fig. 7, Tab. 10).
Species of Nectamia are treated as having gill-raker variable populations because no evidence of consis-
tent color differences could be demonstrated. None of the partially differentiated populations of Nectamia lux-
uria or N. viria have very precisely known distributions based on gill raker counts. Populations with
intermediate gill-raker counts should be expected with better geographic sampling between the lower count
Philippino-Indonesian region and adjacent areas.
Acknowledgments
For the loan of material and the use of museum facilities thanks are extended to many colleagues over the past
decades. I especially recognize the late Ernest A. Lachner with whom I started a portion of this revision in
1974 while at the National Museum of Natural History. The following individuals: C. Lavett Smith (AMNH),
Doug Hoese (AMS), William G. Saul, Eugenia Böhlke (ANSP), David Catania, William N. Eschmeyer and
Tomio Iwamoto (CAS), Oliver Crimmen, Patrick Campbell and Darrell Siebert (BMNH), John E. Randall,
Arnold Y. Suzumoto (BPBM), Robert K. Johnson (FMNH), Karsten Hartel (MCZ), Erling Holm and Richard
Winterbottom (ROM), Uwe Zajonz, Friedhelm Krupp, and Horst Zetzche (SMF), Rob Robins and George
Burgess (UF), Susan L. Jewett, Sandra Raredon, Lisa Palmer, David G. Smith, and Jeffery T. Williams
(USNM), and Ernest D. Estevez, Mote Marine Laboratory aided in curatorial processes. David G. Smith and
Sandra Raredon provided radiographs of several species. G. Duhamel (MNHN) loaned the specimens col-
lected during the expedition of L. de Freycinet for the 1817–1820 voyage around the world by the Uranie and
Physicienne. Dorthea Hubbs took many radiographs of apogonids during 1967–68 for E. A. Lachner. Gerald
R. Allen (WAM), Rudie H. Kuiter, Aquatic Photographics, Robert F. Myers, Coral Graphics, Ukkrit
Satapoomin, Phuket Marine Biological Center, John E. Randall (BPBM) and Jeffery T. Williams provided
color transparencies of some of the species. William N. Eschmeyer and Victor G. Springer (USNM) provided
useful comments concerning nomenclatural treatment. The author was supported by a Smithsonian post-doc-
toral fellowship for an early part of this study. Leonard P. Schultz funds were provided by for several study
trips to the Smithsonian. I thank two reviewers (David Greenfield and anonymous) for their constructive com-
ments.
Zootaxa 1691 © 2008 Magnolia Press · 51
status of Nectamia & 4 new species
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