A revision of the goby genus Gnatholepis Bleeker (Teleostei, Gobiidae, Gobionellinae), with description of a new species

Abstract

Species of the tropical gobiid genus Gnatholepis have been misidentified and confused for many years. A gobionelline, Gnatholepis is the only strictly marine genus of that subfamily. Gnatholepis was reviewed from a morphological viewpoint, as recent publications distinguishing species (morphologically and genetically) have not always been in agreement. The genus is considered to include nine described species, with two distinctive sister-taxa (G. argus and G. gymnocara) and seven less easily distinguished taxa: one species-group formed by G. anjerensis, G. knighti, G. ophthalmotaenia andG. yoshinoi and another species-group of G. cauerensis, G. pascuensis and G. thompsoni. Of these, the G. cauerensis species-group is the most complex, with several subspecies-names available. There is also a distinctive taxon from the Red Sea that represents a new species, described herein.

Full text

Accepted by A. Gill: 18 Sept. 2012; published: 29 Oct. 2012
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Copyright © 2012 · Magnolia Press
Zootaxa 3529: 1–69 (2012)
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ZOOTAXA
A revision of the goby genus Gnatholepis Bleeker
(Teleostei, Gobiidae, Gobionellinae),
with description of a new species
HELEN K. LARSON
1
& DUNCAN J. BUCKLE
2
1
Museum and Art Gallery of the Northern Territory, PO Box 4646, Darwin, NT 0801, Australia. Email: helen.larson@nt.gov.au.
2
Research Institute for the Environment and Livelihoods, Charles Darwin University, Darwin NT 0909, Australia,
Email: Duncan.Buckle@cdu.edu.au.
Magnolia Press
Auckland, New Zealand
3529
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HELEN K. LARSON& DUNCAN J. BUCKLE
A revision of the goby genus Gnatholepis Bleeker (Teleostei, Gobiidae, Gobionellinae), with description
of a new species
(Zootaxa 3529)
69 pp.; 30 cm.
29 Oct. 2012
ISBN 978-1-77557-028-8 (paperback)
ISBN 978-1-77557-029-5 (Online edition)
FIRST PUBLISHED IN 2012 BY
Magnolia Press
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ISSN 1175-5326 (Print edition)
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Gnatholepis Bleeker, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Key to species of Gnatholepis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Gnatholepis anjerensis (Bleeker, 1851) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Gnatholepis argus Larson and Buckle, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Gnatholepis caudimaculata new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Gnatholepis cauerensis (Bleeker, 1853) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Gnatholepis gymnocara Randall and Greenfield, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Gnatholepis knighti Jordan and Evermann, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Gnatholepis ophthalmotaenia (Bleeker, 1854) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Gnatholepis pascuensis Randall and Greenfield, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Gnatholepis thompsoni Jordan, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Gnatholepis yoshinoi Suzuki and Randall, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Incertae sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Abstract
Species of the tropical gobiid genus Gnatholepis have been misidentified and confused for many years. A gobionelline,
Gnatholepis is the only strictly marine genus of that subfamily. Gnatholepis was reviewed from a morphological view
-
point, as recent publications distinguishing species (morphologically and genetically) have not always been in agreement.
The genus is considered to include nine described species, with two distinctive sister-taxa (G. a rg u s and G. g y m n o c a r a )
and seven less easily distinguished taxa: one species-group formed by G. a n j e r e n s i s , G. k n i g h t i , G. o p h t ha l mo t a e ni a and
G. y os h i no i and another species-group of G. cauerensis, G. p a s c u e n s i s and G. thompsoni. Of these, the G. c a u ere n s i s spe
-
cies-group is the most complex, with several subspecies-names available. There is also a distinctive taxon from the Red
Sea that represents a new species, described herein.
Keywords: Gnatholepis, Indo-Pacific, Atlantic, Gobiidae, Gobionellinae, new species
Introduction
The tropical marine gobiid genus Gnatholepis has been misidentified and confused for many years, despite an
abundance of material available for study in museum collections (e.g. see Hoese 1986; Myers 1999; Randall &
Greenfield 2001). There are 18 nominal taxa that belong to the genus. Gnatholepis is unusual in that it is the only
fully marine genus of the subfamily Gobionellinae, species of which are mostly estuarine to fresh water dwelling
(Larson 2001; Thacker 2004a). Members of the genus Gnatholepis are habitat-specific gobies, being found on sand
or silty sand, always near corals.
The genus was originally created by Bleeker (1874), as a subgenus of Stenogobius, with Gobius anjerensis
Bleeker, 1851, as the type species. Gobius anjerensis was described from a Kuhl and van Hasselt illustration of
48 mm specimen (now lost). The illustration shows no colour pattern but it does resemble a Gnatholepis (see
Bleeker 1983), but there is little information that can be obtained from the illustration or the original description
to confirm exactly which species of Gnatholepis the type species actually is. Randall and Greenfield (2001)
recognised the taxonomic problems that could arise from this, and designated a neotype for Gobius anjerensis
(BPBM 26651).
Species presently considered to be Gnatholepis were originally described in either the genus Gobius or in
Gnatholepis (Table 1), although a variety of species with scales on the cheeks and opercles has been placed in the
genus in the past, such as Arcygobius baliurus, Exyrias puntang, Hemigobius mingi and Macrodontogobius wilburi
(Murdy & Hoese 1984; Murdy 1985; Larson 1999; Larson & Wright 2003).
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Most recent publications in which Gnatholepis features, discuss relationships. For example, Birdsong et al.
(1988), in their review of gobioid relationships based on osteology, discussed Gnatholepis within their Gobionellus
Group of the subfamily Gobiinae, Harrison (1989) placed Gnatholepis in the subfamily Gobionellinae (based on
several osteological characters), as did Pezold (1993) and Larson (2001), who both described characters defining
the subfamily. In 2009, Thacker proposed a new classification of gobioids based on DNA sequence data from
several studies, with the Gobionellinae elevated to a new family, Gobionellidae (which also included the
Amblyopinae, Oxudercinae and Sicydiinae), with no supporting characters or synapomorphies provided. This
classification is not followed here (and see Mooi and Gill 2010: 37).
Recent attempts at distinguishing species have been few and not always in agreement (e.g. Hoese 1986; Kuiter
& Tonozuka 2001; Randall & Greenfield 2001; Nakabo 2002). Brito and Miller (2001) included an illustrated
description of Gnatholepis thompsoni, which included a generic diagnosis.
Characters of the genus Gnatholepis were reviewed by Randall and Greenfield (2001), and they also
described a new species from northern Australia (G. g y m n o c a r a ) and four new subspecies of G. c a u e r e n s i s . They
created the neotype for Gobius anjerensis Bleeker, 1851 (mentioned above), but also a neotype for Gnatholepis
davaoensis Seale, 1910, without explanation as to why this was necessary. Additionally, they synonymised
several species, without explanation, with Gnatholepis anjerensis. Randall and Greenfield also synonymised
several other species of Gnatholepis without providing reasons for doing so. For example, they placed Gobius
ophthalmotaenia Bleeker as a synonym of Gnatholepis anjerensis, apparently without examination of type
specimens (syntypes exist and are in good condition). Randall and Greenfield’s work was the first attempt by
anyone to review the genus.
Larson and Wright (2003) briefly discussed some taxa that did not belong in Gnatholepis but had been placed
there by previous workers (such as Gobius baliurus and Gnatholepis volcanus). Other taxa that were originally
described in Gnatholepis but do not belong there are Gnatholepis mingi Herre, 1936 (= Hemigobius mingi; Larson
1999), Gnatholepis hendersoni Herre, 1936 (= Macrodontogobius wilburi; Murdy & Hoese 1984), Gnatholepis
koumansi Herre, 1937 (= a species of Aulopareia, based on examination of the holotype by the senior author) and
Gnatholepis sindonis Snyder, 1909 (= Exyrias puntang; Murdy 1985). The type of Gnatholepis turneri Roxas and
Ablan, 1940, was destroyed in WWII and it is not clear from the description what genus the authors actually had.
However, the senior author’s examination of some Acentrogobius specimens from Batangas fishmarket,
Philippines (CAS 51408 and CAS 52004) found that these may be conspecific with Roxas and Ablan’s specimen,
agreeing well in counts and general appearance. These specimens are presently under study by Zeehan Jaafar of the
National University of Singapore (who is revising Acentrogobius). Two uncertain taxa that may be Gnatholepis are
Eleotris feliceps Blyth, 1860 and Gobius pauper De Vis, 1884 (discussed below).
Thacker (2003) presented a molecular phylogeny of 67 gobioid species, including three Gnatholepis species,
which she identified as G. c a u e re n s i s , G. s c a p u l o s t i g m a and G. t h o m p s o n i . The last two species appeared as a sister-
group in her cladogram (Thacker 2003: fig. 1) and all three Gnatholepis appeared as the sister-group of the
Evorthodus and Ctenogobius sister-group within the gobionelline clade.
Thacker (2004a) discussed the species of Gnatholepis and carried out a cladistic analysis, using DNA data, of
what she considered to be six species. However, the names she assigned to each Gnatholepis species did not
coincide with those used by Randall and Greenfield (2001) and she did not examine type material of G. c a u e re n s i s
or G. s c a p u l o s t i g m a , despite using the name G. s c a p u l o s t i g m a for the species usually referred to by other workers
as G. c a u e re n s i s (e.g. Randall & Greenfield 2001). Thacker also here apparently used the name G. a n j e re n s i s for the
species identified as G. c a u e re n s i s in her 2003 phylogeny paper. She did find that G. a r g u s Larson and Buckle (as
Gnatholepis sp.) was the most plesiomorphic species (Thacker 2004a). There are some problems with the
identification of her material, however. For example, she lists (and illustrates their localities on her Fig. 1) three
species from the Northern Territory, Australia (G. a n j e re n s i s , G. d a v a o e n s i s , G. sp. (= G. a r g u s )). Only one species
exists in this area (G. a rg u s ; Larson & Buckle 2005). Material identified by Thacker as G. a n j e r e ns i s and G.
davaoensis from the Northern Territory are all G. a rg u s , based upon direct examination, by the senior author, of the
same specimens examined by Thacker (Larson & Buckle 2005). It is possible that other specimens in Thacker’s
study had been misidentified. Thacker et al. (2008) presented a similar picture of six species of Gnatholepis, based
on previous work with the addition of specimens of G. c a u e re n s i s (as scapulostigma) from Papua New Guinea.
Thacker (2004b) carried out an analysis of the population structures of two species of Gnatholepis, from Fiji to
the Tuamotus. She identified the fish as G. a n j e r e ns i s and G. s c a p u l o s t i g m a (the latter species is referred to herein
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
as G. c a u e r e n s i s ), and observed that G. a n j e re n s i s , inhabiting shallow lagoonal habitats (0.1–41.3 m depths
(Thacker 2004b)), exhibited more population structuring than did G. c a u e re n s i s , which generally occurs in deeper
reef slope habitats (1.7–89.0 m).
Rocha et al. (2005) presented a scenario, based on mtDNA, that Gnatholepis invaded the Atlantic from the
Indian Ocean about 145,000 years ago, and spread out into the central and eastern Atlantic 100,000 years ago (and
are continuing to invade localities such as the Canary Islands as sea temperatures rise). Rocha et al. considered G.
thompsoni and G. c a u e re n s i s to be sister taxa; they referred to the latter species as G. s c a p u l o s t i g m a .
Larson and Buckle (2005) described a new species of Gnatholepis from northern Australia and commented
briefly on some of the problems created by previous workers and the need for a generic revision of the genus. After
asking many people for specimens and
colour photographs of live and freshly dead Gnatholepis, this study
represents that revision.
TABLE 1. Nominal species of Gnatholepis and their status.
Nominal species Taxonomic allocation Authority
Gobius anjerensis Bleeker, 1851 Gnatholepis anjerensis (Bleeker, 1851) Randall & Greenfield 2001
Gnatholepis argus Larson & Buckle, 2005 Gnatholepis argus Larson & Buckle, 2005 Larson & Buckle, 2005
Gnatholepis calliurus Jordan & Seale, 1905 Arcygobius baliurus (Valenciennes, 1837) Larson & Wright 2003
Gobius capistratus Peters, 1855 Gnatholepis anjerensis (Bleeker, 1851) This work
Gobius cauerensis Bleeker, 1853 Gnatholepis cauerensis (Bleeker, 1853) This work
Gnatholepis cauerensis australis Randall &
Greenfield, 2001
Gnatholepis cauerensis (Bleeker, 1853) This work
Gnatholepis cauerensis hawaiiensis
Randall & Greenfield, 2001
Gnatholepis cauerensis (Bleeker, 1853) This work
Gnatholepis cauerensis pascuensis Randall
& Greenfield, 2001
Gnatholepis pascuensis Randall & Greenfield,
2001
Randall 2009
Gnatholepis corlettei Herre, 1935 Gnatholepis ophthalmotaenia (Bleeker, 1854) This work
Gnatholepis davaoensis Seale, 1910 Gnatholepis ophthalmotaenia (Bleeker, 1854) This work
Gobius deltoides Seale, 1901 Gnatholepis anjerensis (Bleeker, 1851) This work
Eleotris feliceps Blyth, 1860 Possibly Gnatholepis or Asterropteryx From original description
Gnatholepis gemmeus Herre, 1927 Gnatholepis ophthalmotaenia (Bleeker, 1854) This work
Gnatholepis gymnocara Randall &
Greenfield, 2001
Gnatholepis gymnocara Randall & Greenfield,
2001
Randall & Greenfield 2001
Gnatholepis hendersoni Herre, 1936 Macrodontogobius wilburi Herre, 1936 Murdy & Hoese 1984
Gnatholepis inconsequens Whitley, 1958 Gnatholepis cauerensis (Bleeker, 1853) Randall & Greenfield 2001
Gnatholepis knighti Jordan & Evermann,
1903
Gnatholepis anjerensis (Bleeker, 1851) Thacker 2004
Gnatholepis koumansi Herre, 1937 Aulopareia examination of holotype by
senior author
Gnatholepis mingi Herre, 1936 Hemigobius mingi (Herre, 1936) Larson 1999
Gobius ophthalmotaenia Bleeker, 1854 Gnatholepis ophthalmotaenia (Bleeker, 1854) This work
Gobius pauper De Vis, 1884 Possibly Gnatholepis gymnocara This work
Gnatholepis scapulostigma Herre, 1953 Gnatholepis cauerensis (Bleeker, 1853) Randall & Greenfield 2001
Gnatholepis sindonis Snyder, 1909 Exyrias puntang (Bleeker, 1851) Murdy 1985
Gnatholepis thompsoni Jordan, 1904 Gnatholepis thompsoni Jordan, 1904 Thacker 2004
Gnatholepis turneri Roxas & Ablan, 1940 Acentrogobius ? Larson & Wright 2003
Gnatholepis volcanus Herre, 1927 Possibly Exyrias Larson & Wright 2003
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Material and methods
Morphometrics and measurements. Measurements were taken using electronic callipers and dissecting
stereomicroscope. Counts and methods generally follow Hubbs and Lagler (1970), except for transverse scale
counts (TRB), taken by counting the number of scale rows from just before the anal fin origin diagonally
upward and back toward the second dorsal fin base, head length is taken to the upper attachment of the
opercular membrane, body width taken at anus, and depth of body is taken twice: at first dorsal fin origin and
also at anus. First dorsal fin spine lengths were taken, as the shape of the fin was thought to have species-
specific significance, but as the tips of the spines were often curved (this depended upon preservation and
whether the fin was preserved adpressed or partly extended), the spine measurements were interpreted with
some caution (see below in individual species accounts). Predorsal scale counts were made by counting rows,
as usually only the anteriormost two or three scales were on the mid-line of the nape, with other scale rows
crossing at the midline.
Morphometrics are expressed as a percentage of standard length (SL) or head length (HL). In the description,
numbers in parentheses after counts indicate the number of specimens with that count, or the range of counts.
Pterygiophore formula follows Birdsong et al. (1988). Vertebral counts and other osteological information were
obtained by clearing and double-staining and by X-ray. Terminology for lateral canals and sensory pores follows
Larson (2001). Papillae rows are named based on Sanzo (1911).
Abbreviations for institutions referred to are as in Leviton et al. (1985), with the exceptions of: NTM (Museum
and Art Gallery of the Northern Territory, Darwin), PMR (Prirodoslovni Muzej Rijeka, Croatia), RMNH
(Nationaal Natuurhistorisch Museum, Leiden), SAIAB (South African Institute of Aquatic Biodiversity,
Grahamstown) and ZRC (Raffles Museum of Biodiversity Research, Singapore).
In the species accounts provided below, full descriptions of G. a rg u s , G. g y m n o c a r a , G. p a s c u e n s i s and G.
yoshinoi are omitted, as these have recently been adequately described and illustrated in Larson and Buckle (2005),
Randall and Greenfield (2001) and Suzuki and Randall (2009). Diagnoses and remarks are presented for each
species.
Principal co-ordinates analyses. Canonical analysis of principal co-ordinates (CAP) has been used as a
generalized discriminant analysis based on distances (Anderson & Robinson 2003; Anderson & Willis 2003). We
used this to determine the distinctness of assigned Gnatholepis species groups using the morphometric counts and
measurement proportions given in Table 2 (a subset of all data recorded). The analysis was conducted using
PERMANOVA + software (Anderson et al. 2008), based on Euclidean distance. The number of principal
coordinate axes used to discriminate the species groups is the number of axes (m) that resulted in the smallest
cross-validation error (or smallest misclassification error). The misclassification error measures the effectiveness
of the CAP model to distinguish between the species by removing one sample at a time and apply the canonical
model from all the other samples to the excluded sample in order to place it into the canonical space and allocate it
to a particular species (leave-one-out procedure).
The CAP graphic is a constrained ordination and illustrates the best possible separation of species in canonical
space (multidimensions, with samples (represented by each point) closest together being the more similar along the
displayed axes). Vectors have been overlaid onto the CAP graphic to illustrate the direction of influence for the
underlying morphometric counts and proportions. A ‘multiple’ correlation type has been used to calculate vectors
to ensure the displayed direction of influence for each variable takes into consideration the combined influence of
all other variables included in the analysis. The vector length indicates the strength of correlation along the
displayed axes, shorter vectors typically indicate the correlation for that variable is along another axis in canonical
space.
The sub-set of fish specimens included (258 specimens with usable data) in the analysis are detailed in
Appendix 1. The morphometric counts and meristic data used in the analysis are included to enable the CAP
model to be recreated, if desired. A potentially useful feature of the CAP program in PERMANOVA+ is that
new specimens of unknown identity can be added then analysed and assigned into one of the seven species
groups.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
TABLE 2. Morphometric counts and proportions used in canonical analysis of principal co–ordinates to discriminate
between Gnatholepis species.
Results
Gnatholepis Bleeker, 1874
Stenogobius (Gnatholepis) Bleeker, 1874: 318, type species Gobius anjerensis Bleeker, 1851, by original designation and
monotypy.
Gnatholepis—Koumans 1931: 86–87; Goren 1979: 42; Yoshino in Masuda et al. 1984: 251; Hoese 1986: 790; Maugé 1986:
369; Birdsong et al. 1988: 188; Parenti and Thomas 1998: 270; Brito and Miller 2001: 257; Larson 2001: 42; Randall and
Greenfield 2001: 2; Larson and Murdy 2001: 3592; Thacker and Cole 2002: 840; Murdy and Hoese 2003: 1791; Larson
and Wright 2003: 127; Thacker 2004a: 573; Larson and Buckle 2005: 67.
Diagnosis. First dorsal fin spines VI; second dorsal fin rays I,10–12; anal fin rays I,11–12; pectoral fin rays 14–19;
lateral scale count 26–31; transverse scale count backward (TRB) 9–11. Head and body somewhat compressed,
snout short, steep and may be rounded, mouth rather small, subterminal to inferior, with snout often partly
overhanging tips of jaws; lower lip with thin triangular flap or fold ventroposteriorly (greatly reduced in two
species). Eyes set high on head, orbit partly above dorsal profile. Opercle and preopercle covered with scales in all
but two species (no scales present on side of head).
Lateral canal pores on head with anterior nasal pore just anterior to anterior naris, posterior nasal pore beside
each posterior naris, pair of anterior interorbital pores, single posterior interorbital pore, a postorbital pore, a
terminal pore over opercle and anterior and posterior temporal pore in separate posterior portion of oculoscapular
canal over opercle; three preopercular pores present (Fig. 1). Transverse sensory papillae pattern on head, papilla
rows short; with four short, often broken, vertical rows on cheek below eye, row d in two sections, row b may be
absent or reduced; ot row on opercle usually in two sections and os and oi rows short; in some specimens (Fig. 1);
e rows along lower jaw forming two or three short transverse rows near chin.
Anterior eighth to one-third of first gill slit closed by membrane. Inner face of upper limb of first gill arch (to
lesser extent in other arches), covered with low dense papillae which may form clumps or groups; dorsal portion of
arch may have short fleshy protuberances ending in one or several papillae; outer faces of gill filaments covered
with fine papillae, especially along axis of filament (papillae most pronounced on inner face filaments of first arch
Morphometric code Description
Pectoral right Pectoral fin rays, right side
Pectoral left Pectoral fin rays, left side
Lat. scales Count of lateral scales
TRB Transverse scale count backward
Pred. scales Count of predorsal scales along mid-line
HL in SL Proportion of head length in standard length
HD in HL Proportion of head depth in head length
HW in HL Proportion of head width in head length
BDA in SL Proportion of body depth at anal fin origin in standard length
BD D1 in SL Proportion of body depth at first dorsal fin spine in standard length
CPL in SL Proportion of caudal peduncle length in standard length
CPD in SL Proportion of caudal peduncle depth in standard length
Snout in HL Proportion of snout length in head length
Eye in HL Proportion of eye length in head length
Jaw in HL Proportion of jaw length in head length
Preorb. in HL Proportion of preorbital width in head length
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and on both faces of filaments of remaining arches). Tongue short, fleshy, bilobed. Anterior naris in short tube, may
have small flap posteriorly; posterior naris low, rounded. Pectoral girdle (edge of cleithrum) smooth, with no fleshy
knobs or lobes. Pelvic fins fused, oval; margin of fraenum fimbriate, fringed with many fine pointed lobes or thin
lappets. Genital papilla in males small, narrow and slender, narrowing to pointed tip; papilla in females short,
fleshy and rounded.
FIGURE 1. Sensory pores and papillae in Gnatholepis anjerensis, SAIAB 2112, Mauritius, 46 mm SL male. Scales omitted.
Characteristic colour pattern with black mark atop eyeball, dusky to black line from eye crossing cheek to
rictus, dark and/or light spot or blotch above pectoral base; body pale with variable pattern of rows of dark lateral
blotches, spots and fine lines.
Dorsal pterygiophore pattern 3-12210; 26 vertebrae (10 precaudal and 16 caudal), rarely 10+15 or 10+17;
usually two epurals (usually one in G. o p h t h a l m o t a e n i a ), two anal fin pterygiophores present before first caudal
haemal spine; neural spines of first few vertebrae slender, pointed; fifth ceratobranchials narrowly triangular,
interdigitated posteriorly along median line and with high flange ventrally, dorsally covered with long slender
teeth; metapterygoid short, relatively small, with short triangular dorsal process, not contacting or forming bridge
to quadrate; palatine stout, broadened anteriorly, not reaching quadrate; pterygoid small, relatively slender, less
than half the length of palatine, in contact with quadrate; quadrate somewhat forked; three or four ossified gill-
rakers; scapula unossified; first epineural inserting on parapophysis of first vertebra.
Remarks. Having the fifth ceratobranchials medially interdigitated (especially posteriorly) is a character that
Gnatholepis shares to some degree with Stenogobius (in which the fifth ceratobranchials are partly interdigitated in
a few species; character not widely reviewed) (Larson 2001). In Evorthodus lyricus (an Atlantic gobionelline) these
bones are joined but not interdigitated (Parenti & Thomas 1998; HKL pers. observ.). Most gobioids have these
bones separate.
Gnatholepis species have the frenum edged with a fine membranous fringe, unlike most gobies, which have a
smooth even edge to the frenum. The fimbriate frenum is a feature shared by several gobionelline genera, such as
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Oxyurichthys, Oligolepis and Gobionellus (Pezold & Larson in prep.). In Stenogobius the frenum is smooth to
finely crenulate (HKL pers. observ.).
Watson (1992) illustrated the small fleshy papillae along the axes of each gill filament in Awaous and
considered that the feature was unique to that genus. The papillae covering most of the gill filaments and
undersides of gill arches in Gnatholepis greatly resemble those present in species of Awaous. Stenogobius
laterisquamatus also possesses fine papillae along the gill filaments, though papillae are more sparse and absent
from the gill arches (HKL pers. observ.). We found that these fleshy papillae are present along the gill arches and
gill filaments of all species of Gnatholepis. This feature bears further observation.
All but two species of Gnatholepis possess a small triangular flap or fold in the end of the lower lip near the
rictus; its function is unknown. The two small species Gnatholepis argus and G. g y m n o c a r a lack this structure, as
do most gobioids.
Gnatholepis has the dorsal portion of the gill arch with one or more short fleshy protuberances extending
into the oral cavity and ending in one or several papillae. They greatly resemble those fleshy lobes present in
sand-dwelling gobiines such as Va le n ci e n n e a and Amblygobius (Hoese & Allen 1977: fig. 4). The condition in
Awaous is similar, but it has a broad fleshy pad covered with small papillae (similar to those covering the gill
filaments).
Gnatholepis species have characteristic variably developed black blotches or lines over the dorsal part of the
eye. The shape and arrangement of these markings often seem to be species-specific (see species accounts).
Additionally, the upper half (at least) of the eyeball is known to fluoresce under red light in at least one species of
Gnatholepis (Michiels et al. 2008); this feature has not yet been widely surveyed. The black marking on the dorsal
surface of the eye may be part of inter-species communications.
Results of CAP analysis of Gnatholepis. The canonical analysis of principal co-ordinates (CAP) on seven
Gnatholepis species (G. a n j e r e n s i s , G. c a u d i m a c u l a t a n. sp., G. c a u e re n s i s , G. k n i g h t i , G. o p t h a l m o t a e n i a , G.
pascuensis and G. t h o m p s o n i ; Table 3) is illustrated in Figure 2. CAP axes 1 and 2 explain 62% of the total
variation within the 13 (m = 13) canonical axes required to best discriminate between the species. In Figure 2, G.
anjerensis occupies the uppermost region of the CAP plot separating from other species along axis 2 by
differences, primarily, in the proportion of: head length to standard length, preorbital width in head length and jaw
length in head length. Gnatholepis thompsoni, G. c a u e re n s i s and G. p a s c u e n s i s occupy the right-hand side
separating most along axis 1 primarily by increased pectoral ray counts and lateral scale counts. Gnatholepis
caudimaculata, G. o p h t h a l m o t a e n i a and G. k n i g h t i occupy the left-hand side separating along axis 1 primarily
influenced by greater proportions of head and eye widths in the head length. Although the seven species cluster
into different areas in ordination space, there are no distinct separations illustrated by the CAP plot, although
separation of some species could occur along other canonical axes given that 38% of the canonical variation is not
illustrated in Figure 2.
The leave-one-out allocation success is based on the entire 13 canonical axes and showed 64% (183/285)
overall correct allocation. However, the allocation success of individual species varies considerably (Table 3). The
only distinct species shown is G. p a s c u e n s i s which achieved 100% correct allocation of the five samples.
Gnatholepis thompsoni had a good classification rate of 81.6% whilst G. o p t h a l m o t a e n i a and G. a n j e re n s i s had
reasonable classification success (73.8% and 71.2%, respectively). Gnatholepis knighti, G. c a u e re n s i s and G.
caudimaculata n. sp. had lower classification success rates (61.9%, 49.4% and 48% respectively) (Table 3). These
results support the CAP illustration (Figure 2) that Gnatholepis species have subtle differences in their
morphometric counts and proportions and that the effectiveness of the selected variables to discriminate species
varies. Most importantly, meristic counts and morphometric proportions do not provide clear taxonomic
identification, with the exception of G. p a s c u e n s i s (a very small sample size).
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FIGURE 2. Canonical analysis of principal co-ordinates plot for the seven Gnatholepis species with vectors overlaid. CAP
axes 1 and 2 explain 62% of the variation from 13 canonical axes.
Several Gnatholepis species have distinctive features and/or geographic distributions that can be used to
distinguish, or isolate, that species from others. These species include G. ca u d i m a c u l a t a n. sp., which has a
distinctive vertical oval blackish to dark grey blotch across the caudal fin base and is known from the Red Sea and
Persian Gulf; G. o p t h a l m o t a e n i a has distinctive black spots on the anal and dorsal fins and breast scales that reach
forward to the rear margin of the preopercle (with an Indo-West Pacific distribution); and G. p a s c u e n s i s has high
pectoral fin ray counts (18–19) with a distinctive pattern of two rows of five elongate dense black blotches on the
side of the body and is known only from Rapa. Removal of these three distinguishable species from the CAP
analysis reduces unnecessary misclassification of the other species which do not share these distinctive features
(for example 18, 13 and 6 specimens were incorrectly assigned as G. c a u d i m a c u l a t a n. sp., G. o p t h a l m o t a e n i a and
G. p a s c u e n s i s respectively (see Table 3)).
Re-analysis of the remaining four species (G. a n j e re n s i s , G. t h o m p s o n i , G. c a u e re n s i s and G. knighti) using
CAP results in a small improvement to the overall misclassification success (69% or 144/209, Table 4; Fig. 3).
Gnatholepis cauerensis and G. knighti have the lowest classification success (57.5% and 66.7% respectively)
(Table 4). Gnatholepis cauerensis is most commonly misclassified with G. t h o m p s o n i (Table 4: 18 samples; G.
thompsoni is restricted to the Atlantic) and G. k n i g h t i (12 samples; restricted to Hawaii), while G. k n i g h t i is most
commonly misclassified with G. c a u e r e n s i s (5 samples). Gnatholepis thompsoni and G. a n j e re n s i s had quite good
classification success (81.6% and 76.9% respectively).
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
TABLE 3. Allocation success results for all seven identified species of Gnatholepis using a canonical analysis of
Principal Co-ordinates Analysis (CAP) based on 16 morphometric counts and proportions. Values in bold show the
number of specimens for each species that have been correctly allocated.
TABLE 4. Allocation success results for four identified species of Gnatholepis (not always easily distinguishable by
pigment colour) using a canonical analysis of Principal Co-ordinates Analysis (CAP) based on 16 morphometric counts
and proportions. Values in bold show the number of specimens for each species that have been correctly allocated.
Original group
G. anjerensis
G. caudimaculata n. sp.
G. cauerensis
G. knighti
G. ophthalmotaenia
G. pascuensis
G. thompsoni
Total # of specimens for
each species
Total # of specimens
missallocated to other
species
% correctly allocated to
each species
G. anjerensis 37 4243 0 2521571.15%
G. caudimaculata n. sp. 4 14 128 0 0291548.28%
G. cauerensis 5743 10 1 5 16 87 44 49.43%
G. knighti 32213 10021 8 61.91%
G. ophthalmotaenia 241331 01421173.81%
G. pascuensis 00000 5 05 0100.00%
G. thompsoni 21500 140 49 9 81.63%
Total # of specimens that
are incorrectly assigned by
CAP to each species
16 18 11 19 13 6 19
Total # of specimens used = 285
Total # specimens correctly assigned = 183 (64.21%)
Total # specimens misclassified = 102 (35.79%)
Original group
G. a n j e r e n s i s
G. c a u e re n s i s
G. k n i g h t i
G. t h o m p s o n i
Total # of specimens for
each species
Total # of specimens
missallocated to other species
% correctly allocated
to each species
G. anjerensis 40 282 52 12 76.92%
G. cauerensis 7 50 12 18 87 37 57.47%
G. knighti 2514 0217 66.67%
G.thompsoni 36040 49 9 81.63%
Total # of specimens that are
incorrectly assigned by CAP to
each species
12 13 20 20
Total # of specimens used = 209
Total # specimens correctly assigned = 144 (68.9%)
Total # specimens misclassified = 65 (31.1%)
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FIGURE 3. Canonical analysis of principal co-ordinates plot for the four Gnatholepis species without reliably distinctive
colour pattern, with vectors overlaid. CAP axes 1 and 2 explain 64% of the variation from 14 canonical axes.
Analysis of meristics and proportions using the CAP analyses procedure above shows that the seven
Gnatholepis species analysed differed subtly in body shape and/or scale or fin ray counts, with only one species, G.
pascuensis, distinctly different. Characteristic body/fin pigment and geographic location were identified by the
authors to distinguish three species: G. c au d i m a c u l a t a n. sp., G. o p t h a l m o t a e n i a and G. p a s c u e n s i s . Four species (G.
anjerensis, G. thompsoni, G. c a u e re n s i s and G. knighti) remain problematic for reliable identification using meristic
counts, morphometric proportions, body and fin pigment and geographic location. The authors have used all the
features above (as well as the 'gestalt' of the specimens as an identification tool) when sorting specimens of
Gnatholepis to species, as a number of the museum lots examined contained more than one species. While current
information does not reliably and easily distinguish all species, it is possible that a deeper understanding of live
specimen colour and genetic differences in combination with taxonomic information may provide better
identification of species within Gnatholepis.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Key to species of Gnatholepis
(may not always work, especially with small juveniles or faded preserved specimens)
1. No scales present on cheek or opercle; predorsal midline naked (few scales may cross nape close behind eyes); nape scales
always cycloid; no enlarged canine teeth in lower jaw. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Opercle and cheek scaled; predorsal with scales crossing midline; nape scales ctenoid and/or cycloid; enlarged canine or
recurved teeth present in lower jaw. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Second dorsal fin I,10, rarely I,11; anal rays I,11; no scales on pectoral fin base; males with small black spots on unpaired fins,
and posteriormost 3–4 mid-lateral blotches darker than anterior spots (northern Australia) . . . . . . . . . . . . . . . . . . . . . . G. argus
– Second dorsal fin I,11, rarely I,10; anal rays I,12; few embedded scales on pectoral fin base; males without dense black spots
on fins and no large dark blotches on side of body (Queensland, Australia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .G. gymnocara
3. Anal fin with 1–4 irregular rows of oval black spots (fin also with red, blue and yellow markings when alive), and first dorsal
fin with 2–4 dense black spots along fin base in both sexes, caudal fin in males with staggered rows of small dense black spots;
each scale on anterodorsal half of body with black or blackish spot; breast scales reaching forward anterior to rear margin of
preopercle or further forward to below mid-eye; nape scales often mostly ctenoid (Cocos-Keeling, Philippines, Taiwan, south-
ern Japan, Indonesia, New Guinea to Vanuatu) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. ophthalmotaenia
– Anal fin without 1–4 rows of dense black spots, although fin may be heavily pigmented and irregular dusky spots may be
formed, first dorsal fin without 3–4 dense black spots along base, caudal fin without rows of dense black spots (may be dusky-
speckled or with brownish spots or streaks); scales on anterodorsal half of body may or may not bear dusky spots; breast scales
reaching forward to rear margin of preopercle; nape scales ctenoid or cycloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. First dorsal fin rather tall, usually pointed, with third spine longest; black blotch usually present on upper part of first dorsal fin
behind first and third spines; pelvic fins banded with light and dark pigment; pectoral rays 15 (data from only 3 specimens)
(Malaysia; Indonesia; Japan). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. yoshinoi
– First dorsal fin convex to almost square; no black blotch present on upper part of first dorsal fin; pelvic fins without dusky
cross-bands; pectoral rays 15–19 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Predorsal scales mostly ctenoid, often extending up to close behind eyes, may be cycloid toward midline and anteriorly; pecto-
ral fin with many rows of fine dark and white speckles; when live, nape and body covered with irregular rows of small blackish
to reddish spots, with five to six diffuse dusky blotches along the lower mid-side of the body; pectoral fin with rows of fine
dark and white speckles; fifth D1 spine as long or nearly as long as 3
rd
–4
th
spines, giving fin rather square appearance when
extended; head and body with stocky appearance (Indo-Pacific, from South Africa to Society Islands) . . . . . . . . G. anjerensis
– Predorsal scales always cycloid in midline, may be some ctenoid scales above opercle; pectoral fin may have dusky lines at
base but not speckled with fine dark spots, if fine dark and light spots present then anal fin with dark streaks or distinct spots on
membrane; when live, colour not as above; pectoral fin translucent and may have whitish or dusky speckling (fine dark and
white speckles together in one species, G. k n i g h t i from Hawaii); fifth D1 spine usually shorter than 3
rd
–4
th
spines, giving fin
rounded appearance when extended; body appears relatively slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Pectoral fin covered with many rows of fine dark and white speckles; transverse black mark on upper part of eye usually split
in two thin lines; anal fin plain dusky, often with one to several blackish rounded to oval spots posteriorly, or with dark streaks
following membrane (Hawaii; Line Islands) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. knighti
– Pectoral fin may have dusky lines at base but without fine dark and light speckles; transverse black mark on upper part of eye
may be narrow or broad but not split; anal fin plain, without distinct dark spots or streaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. 18–19 pectoral fin rays; distinctive pattern of two rows of five elongate dense black blotches on side of body, anteriormost
three blotches largest, joined by faint broken yellowish brown lines in life, a very broad black cheek-bar and a broad black eye-
bar (both as wide as pupil) crossing the interorbital space (Easter Island). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pascuensis
– 15–17 pectoral fin rays; colour pattern not as above, with black cheek-bar and black marking on top of eye variable in intensity
and width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Distinctive vertical oval blackish to dark grey blotch across caudal fin base, oblique black curved or comma-shaped line on top
of eye and body whitish with finely speckled dorsum; when alive, two broad ladder-like orange to reddish stripes along body;
7–9 predorsal scales, usually 8 (Red Sea to Persian Gulf) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. caudimaculata n. sp.
– No distinctive vertical oval blackish to dark grey blotch across caudal fin base; black line on top of eye not curved or comma-
shaped but relatively horizontal and may join its counterpart across interorbital space; 7–12 predorsal scales, usually 9–10. . . . . 9
9. Predorsal scale count 7–12, modally 9, always cycloid in midline, may be some ctenoid scales above opercle; pectoral rays
14–19, modally 16; transverse black line on upper part of eye over centre to rear half of pupil, may partly cross mid-interorbital
space, line joins mostly vertical to curved black line from cheek to past end of jaw; dorsal half of body with 2–3 rows of short
lines or rows of spots (if lower half of body with rows of spots, then these paler than dorsal rows); fifth D1 spine shorter than
3
rd
–4
th
spines, giving fin rounded appearance when extended (Indo-Pacific, from South Africa to Pitcairn Island) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. cauerensis
–. Predorsal scale count 9–11, modally 10; all predorsal scales cycloid; pectoral rays 16–18, modally 17; transverse black line on
upper part of eye joining the somewhat oblique to curved black line crossing cheek and ending well behind end of jaw; body
covered with up to 8 rows of dark brown spots; in males, 3
rd
–4
th
D1 spines longest, fin tending to have square to rectangular
appearance when extended (Atlantic, from Florida to Canary Islands) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. thompsoni
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TABLE 5. Frequency distribution of fin ray counts in Gnatholepis species (G. y o s h i n o i not included). Data for G.
gymnocara is from Randall and Greenfield (2001) and pectoral ray data for G. p a s c u e n s i s is combined with that in
Randall (2009); for both these species, the pectoral ray counts are halved (as these authors included both sides in their
tables). Data for G. c a u e r e n s i s is split as per Randall (2009) species for comparison purposes.
Species Second dorsal rays Anal rays Pectoral rays (right side whenever possible)
9101112 9101112 14 15 16 17 18 19
G. a n j e r e n s i s –462– –1602 – 11 42 12 – –
G. a r g u s 6605– –8575 – –15505–
G. c a u d i m a c u l a t a n. sp. – 5 31 – – 3 33 – – 9 26 1 – –
G. c a u e re n s i s (Hawaii) – – 14 – – 1 13 – – – 1 14 1 –
G. c a u e re n s i s (Rapa) – 2 17 – – – 11 – 1 1 8 4 6 –
G. c a u e re n s i s (the rest) – 4 78 – – 3 86 2 1 7 40 22 11 1
G. g y m n o c a r a –137– –236– – 111215–
G. k n i g h t i –5213 –424– – 3186––
G. o p h t h a l m o t a e n i a 1153– –353– – 5445––
G. p a s c u e n s i s ––6– 1–5– – – – – 5 5
G. t h o m p s o n i –2541 11551 – –13369–
TABLE 6. Frequency distribution of predorsal scale counts counts in Gnatholepis (G. a r g u s , G. g y m n o c a r a and G.
yoshinoi not included).
Species 6 7 8 9 10 11 12 13
G. a n j e r e n s i s –11323206–1
G. c a u d i m a c u l a t a n. sp. 1 4 20 8 ––––
G. c a u e re n s i s –434541641–
G. k n i g h t i –19124–––
G. o p h t h a l m o t a e n i a 1230157–––
G. p a s c u e n s i s ––231–––
G. t h o m p s o n i –––202991–
TABLE 7. Frequency distribution of transverse backward scale counts in Gnatholepis (G. g y m n oc a r a and G. y o s h in o i not
included).
Species 7 8 9 10 11 12
G. a n j e r e n s i s –– 327308
G. a r g u s 1154014– –
G. c a u d i m a c u l a t a n. sp. – 1 2 22 7 –
G. c a u e re n s i s – – 11 70 28 2
G. k n i g h t i –1 311103
G. o p h t h a l m o t a e n i a –– 103014–
G. p a s c u e n s i s – – – 5 ––
G. t h o m p s o n i –– 226273
TABLE 8. Frequency distribution of lateral scale counts in Gnatholepis species (G. g y m n o c a r a and G. y o s h i n o i not
included).
Species 24 25 26 27 28 29 30
G. a n j e r e n s i s –42626101 –
G. a r g u s 1204494– –
G. c a u d i m a c u l a t a n. sp. – 3 21 7 3 – –
G. c a u e re n s i s 223056205 –
G. k n i g h t i –14 1561 –
G. o p h t h a l m o t a e n i a 110211642 1
G. p a s c u e n s i s ––– 6–– –
G. t h o m p s o n i ––2725271 –
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Gnatholepis anjerensis (Bleeker, 1851)
(Figs 1, 4–7, 11B; Tables 5–9)
Gobius anjerensis Bleeker, 1851: 251, pl. 1, fig. 11 (Anjer, Java Boenakeng, off Ujung Padang, Sulawesi, Indonesia).
Gobius capistratus Peters, 1855a: 251 and 1855b: 443 (Ibo, Mozambique).—Klunzinger 1871: 476–477 (Red Sea); Jatzow and
Lenz 1898: 507 (Zanzibar).
Gobius deltoides Seale, 1901: 125 (Agaña, Guam, Mariana Islands).—Koumans 1940: 156–157; Böhlke 1984: 105.
Gnatholepis deltoides—Yoshino in Masuda et al. 1984: 252 (Yaeyama Islands); Larson and Murdy 2001: 3601; Kuiter and
Tonozuka 2001: 672, figs A–C (Bali); Nakabo 2002: 1212.
Gnatholepis anjerensis—Yoshino i n Masu da et al. 1984: 251 (Ryukyu Islands); Wass 1984: 28 (Samoa); Randall and Randall
1987: 309 (Enewatak Atoll); Randall and Goren 1993: 13 (Maldives); Goren and Dor 1994: 64 (Red Sea); Allen and
Smith–Vaniz 1994: 16 (Cocos (Keeling)); Randall 1995: 336 (Oman); Myers 1999: 255, plate 161D (Micronesia); Fricke
1999: 512 (Mascarene Islands); Larson in Randall and Lim 2000: 637 (South China Sea region); Hutchins 2001: 43
(Western Australia); Larson and Murdy 2001: 3601; Randall and Greenfield 2001: 3; Kuiter and Tonozuka 2001: 671, figs
C, D (Bali, Indonesia); Nakabo 2002: 1212; Hayashi and Shiratori 2003: 96, fig. 176; Allen and Adrim 2003: 58 (Papua to
Sumatra); Manilo and Bogorodsky 2003: S118; Myers and Donaldson 2003: 638 (Marianas); Senou et al. 2004: 244–245;
Randall et al. 2004: 27 (Tonga); Heemstra et al. 2004: 3329 (Rodrigues); Hoese and Larson 2006: 1654; Williams et al.
2006: 259 (Wallis Islands); Kimura et al. 2009: 276 (west coast of Thailand); Fricke et al. 2009: 101.
Acentrogobius cauerensis—Dor 1984: 237 (Red Sea).
Gnatholepis cauerensis—(in part) Maugé 1986: 369 (Transkei northwards); Winterbottom and Emery, 1986: 35 (Chagos
Archipelago); Myers 1991: 238, plate 124A (Guam); Kulbicki and Williams 1997: 22 (Ouvéa Atoll, New Caledonia).
Gnatholepis sp.—Randall et al. 1993: 387 (Midway Atoll).
Gnatholepis sp.1—Kuiter and Tonozuka 2001: 671, fig. (Mabul, Malaysia).
Diagnosis. A large stocky Gnatholepis (up to 84 mm SL) with ctenoid scales on head, predorsal midline and
pectoral fin base, distinct flap present at end of lower lip; at least three dark spots along first spine of first and
second dorsal fins, followed by at least three rows of dark spots or short streaks, pectoral fin with rows of fine dark
and white speckles; second dorsal and anal fin rays nearly always I,11; pectoral rays 14–17, modally 16; lateral
scales 25–29, modally 27; predorsal scales 7–13, usually 9–10, mostly ctenoid, may be cycloid anteriorly or toward
midline.
Material examined. SOUTH AFRICA: SAIAB 2576, 1(84), pool at Thompson’s Bay, Kwa-Zulu Natal, R.
van Elst, 30 May 1972. MOZAMBIQUE: SAIAB 19636, 3(41–52), Inhaca Island, 5 August 1948; SAIAB 16933,
Naca, 27 August 1956; SAM 28944, 5(26–50.6), Ilha Magaruque, S of Bazaruto, B. Kensley and party, 24 March
1973. KENYA: NTM S.13965-005, 1(45.5), Gazi Bay, H. Coene, July–August 1993. MADAGASCAR: USNM
260667, 12(31–51), Tulear [Toliara] Harbour, 11 August 1964; AMS I.28108-038, 4(18.5–44), Andilans Beach,
Nosy Be, 20 September 2005. RODRIGUES: SAIAB 70332, 1(51.5), Yacht Casse, P. Heemstra and party, 21
October 2001. MAURITIUS: SAIAB 2112, 5(26–51), La Morne Brabant, THF-SA-28,7 March 1971; SAIAB
2036, 3(39–44), La Morne Brabant, 6 March 1971; SAIAB 2317, 4(37–45.5), 2 miles N of Trou d’Eau Douce, T.H.
Fraser, 27 March 1971; SAIAB 57877, 3, S of Pointe Petite Riviere, 22 April 1995; SAIAB 57889, 2(42–46),
tidepools at end Avenue Victory, Albion, 25 April 1995. EGYPT: USNM 313779, 1(43), bay on NW end of Giftun
Al Kebir Island, Hurghada, Strait of Jubal, Egypt, H.A. Fehlmann and party, 3 January 1965. ISRAEL: AMS
I.21874-001, 2(35.5–39), Shurat el Mankta, Gulf of Aqaba, L. Fishelson, 15 September 1967; MNHN 1977-778,
1(45), Gulf of Aqaba, M.L. Bauchot and J.E. Randall. YEMEN: USNM 327464, 6(31–45.5), Zubair Island,
Yemen, E. Clark, 30 September 1967. INDONESIA: Neotype of Gobius anjerensis, BPBM 26651, 31 mm SL
male, Bunaken Island, off Manado, Sulawesi, J.E. Randall and party, 28 August 1978; NSMT-P.64544, 3(26–27.5),
Baguala Bay, Ambon, K. Shibukawa, 1 December 1998; NSMT-P.63406, 1(20.5), Lembeh Island, Bitung,
Sulawesi, 13 July 2000; USNM 211025, 19(19–34), bay off Tuhaka, Ihamahu, Pulau Saparua, Moluccas, V.G.
Springer and party, 18 January 1973. GUAM: AMS I.40826-001, 1(44), Tumon Bay, H. Larson, 4 May 1969;
Syntypes of Gobius deltoides, ANSP 84134, 10(28–43), Agana, A. Seale, June 1900; BPBM 267, 4(26–44.5),
Agana, A. Seale, 2 June 1900; USNM 109399, 1(39), A. Seale, 2 June 1900. REPUBLIC OF MICRONESIA:
USNM 224767, 4(28–35), N coast Ponape, Caroline Islands, 4 September 1980. AUSTRALIA: AMS I.28999-010,
2(31.5–36.5), E end of Pulao Kambing, Cocos-Keeling Islands, G.R. Allen, 12 February 1989; NTM S.11973-037,
2(32–37.5), lagoon entrance by West Islet, Ashmore Reef, Timor Sea, H. Larson, 26 July 1986; NTM S.11983-037,
1(33.5), reef flat on W side West Islet, Ashmore Reef, Timor Sea, H. Larson, 29 July 1986; NTM S.11983-002,
1(38), reef flat on W side West Islet, Ashmore Reef, Timor Sea, H. Larson, 29 July 1986; ex-AMS I.20757-072,
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1(40.5), W end Raine Island, Queensland, AIMS-AMS survey, 13 February 1979; AMS I.35852-003, 3(45–60),
first lagoon, north channel, One Tree Island, J. Caley, 7 April 1990; ex-USNM 308212, 2(54–56), reef flat on SE
side One Tree Island, Queensland, V.G. Springer and party, 27 November 1966. SOLOMON ISLANDS: USNM
358895, 1(37), Santa Cruz Island, 22 September 1998. FIJI: USNM 243060, 4(37–49), main harbour on W side
Matuku Island, Lau Group, 23 April 1982. FRENCH POLYNESIA: AMS I.21874-001, 2(35–38), mouth of creek
at head of Opunohu Bay, Moorea, R. Galzin, 15 October 1990; AMS I.30940-006, 8(16–37), mouth of creek at
head of Opunohu Bay, Moorea, R. Galzin, October 1990; CAS 51530, 6(40.5–51), almost-enclosed lagoon,
Taravao, Tahiti, J.E. Randall, 21 April 1956.
Other material; no data taken. SOUTH AFRICA: SAM 25625, 1, Inhaca, B. Kensley and party.
MOZAMBIQUE: SAIAB 16943, 2, Ibo Island, Querimba Archipelago; SAIAB 57255, 4, Querimba Archipelago;
SAIAB 13674, 1, Querimba Archipelago; SAIAB 16928, 2, Pinda Island; SAIAB 16934, 1, Pinda Island. EGYPT:
BPBM 18334, 4, Ras Mohammed, S end Sinai Peninsula; BPBM 19840, 10, beach N of Nuweiba el-Muzeini, Gulf
of Aqaba. SUDAN: BPBM 19760, 1, N of Port Sudan, Red Sea; BPBM 19750, 4, Suakin Harbour, Red Sea.
SAUDI ARABIA: BPBM 30405, 1, about 5 miles N of Marifa, Persian Gulf. MADAGASCAR: USNM 327542, 3.
OMAN: BPBM 36143, 1, Masirah Island; BPBM 34406, 1, Hoone’s Bay, near Mirbat; BPBM 36032, 10, Sawda
Island. ALDABRA: SAIAB 16930, 3; SAIAB 16940, 2; USNM 327760, 60, Aldabra Atoll. SEYCHELLES:
USNM 308213, 46, Farquhar Atoll. PHILIPPINES: AMS I.21931-005, 6, Lapu Lapu market, Cebu, Mactan Island.
INDONESIA: NTM S.15420-009, 1, Pahepa, Sangir Island; AMS I.23679-002, 2, Denpasar, Bali. THAILAND:
NSMT-P.61750, 1, Ko Lon, Phuket. MARIANA ISLANDS: NTM S.14712-001, 2, Sanvitores Beach, Guam.
MICRONESIA: USNM 224763, 59, Ponape, Caroline Islands. MARSHALL ISLANDS: USNM 368807, 5.
VA N U AT U : A M S I B . 3 6 0 7 , 1 , Vi l a , N e w H e b r i d e s . S O C I ETY ISLANDS: MNHN 1984-168, 5, Tiahura, Moorea;
CAS 53895, 2, Moorea; CAS 58267, 2, Moorea; CAS 58268, 2, Moorea; CAS 53896, 2, Papeete Harbour, Tahiti.
Description. Based on 66 specimens, 26–84 mm SL. An asterisk indicates the counts of the neotype of Gobius
anjerensis.
First dorsal VI*; second dorsal I,10–I,11* (mean I,10.9); anal I,10–12 (mean I,11.0*), pectoral rays 14–17
(mean 16.0*), segmented caudal rays nearly always 17*; caudal ray pattern nearly always 9/8*; branched caudal
rays 6/6 to 8/7 (mean 7/6*); lateral scale count 25–29 (mean 26.6, 26 in neotype); TRB 9*–12 (mean 10.4);
predorsal scales 7*–13 (mean 9.3); circumpeduncular scales 11–13 (mean 12.1*). Gill rakers on outer face of first
arch 1–2 + 3–4 (in 6, modally 1+4).
Body compressed, width at anus 11.9–28.1% (mean 15.2%) of SL. Body rather stocky in appearance, body
depth at anus 19.6–26.6% (mean 23.3%) of SL, body depth at first dorsal fin origin 14.3–27.7% (mean 23.3%) of
SL. Head compressed, broader ventrally, slightly deeper than wide, HL 26.6–33.3% (mean 29.3%) of SL; head
depth at posterior preopercular margin 62.0–83.7% (mean 71.2%) of HL; head width at posterior preopercular
margin 53.7–82.2% (mean 68.5%) of HL; head profile bluntly pointed; nape slightly convex behind eyes. Mouth
subterminal to nearly terminal, slightly oblique; jaws generally reaching to below anterior margin of eye; upper jaw
length 32.0–41.5% (mean 35.9%) of HL. Upper lip smooth, narrower than lower, lower lip papillose close to teeth,
with twist or fold posteriorly, forming triangular flap, lip interrupted at chin. Eye moderate to relatively small in
large specimens, dorsolateral, 22.7–30.5% (mean 27.1%) of HL; preorbital width 18.2–27.8% (mean 22.3%) of
HL. Snout bluntly pointed, 22.4–44.4% (mean 34.6%) of HL; posterior naris round to almost triangular, close to
anterior margin of eye; anterior naris in short tube, higher on posterior margin of eye, about level with middle of
eye or somewhat ventral to it. Interorbital narrow, 4.7–11.3% (mean 7.9%) of HL. Caudal peduncle compressed,
length 12.9–18.6% (mean 16.1%) of SL; caudal peduncle depth 10.7–13.8% (mean 11.8%) of SL.
First dorsal fin rounded to square, with no spines greatly elongate; third or fourth spine usually longest; when
adpressed, spine tips reaching to first to third element of second dorsal fin. Third dorsal spine modally longest,
15.4–19.6% (mean 17.6%) of SL; fourth dorsal spine length 15.0–19.8% (mean 17.7%) of SL. Second dorsal fin as
tall as first dorsal fin, rays longer anteriorly than posteriorly, fin pointed to slightly rounded posteriorly. Anal fin
lower than second dorsal fin, anteriormost rays shorter than posterior few rays; fin pointed to slightly rounded
posteriorly. Second dorsal and anal fin rays, when adpressed, usually reaching caudal fin in adults. Pectoral fin
somewhat pointed, central rays longest, 23.5–28.7% (mean 26.0%) of SL; fin reaching back to above first few anal
fin rays. Pelvic fins fused, frenum with distinctive finely fimbriate margin, fins round to somewhat oval, reaching
to first few anal fin rays, 22.1–31.7% (mean 27.4%) in SL. Caudal fin moderate, oval to rounded, 25.1–36.1%
(mean 30.9%) of SL.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Gill opening restricted, extending anteriorly to lower edge of pectoral base or to just under opercle. One or two
slender gill rakers on outer face of first arch, closest to angle of arch, remainder of rakers very short, pointed;
pointed papillae present along inner face of first arch; outer rakers on second gill arch consisting of two series of
papillae, one low and one of slightly pointed papillae; outer rakers on remaining arches low, broad-based. Inner
face of upper limb of first gill arch, and to lesser extent, upper limbs of other arches, covered with low dense fleshy
papillae which may form clumps or groups; dorsal portion of arch may have short fleshy protuberances ending in
one or several papillae. About one-third of first gill arch bound by membrane to inner face of opercle. Tongue
short, tip bilobed to concave.
Teeth in upper jaw in two to three rows across front and one row at side of jaw, outermost row teeth largest,
curved and pointed, largest teeth at front of jaw on either side of symphysis; innermost row teeth very small, sharp
and evenly sized. Teeth in lower jaw in two to three rows, arranged similarly to upper jaw but outer row teeth may
be smaller (in females); posteriormost one to three outer row teeth enlarged and somewhat recurved in males.
Ctenoid predorsal scales on nape of variable extent, from just over opercle to close up behind eyes; nape
midline scales may be partly cycloid from behind eyes to rear margin of preopercle, nape midline never entirely
with cycloid scales (Fig. 4). Opercle covered with scales, cycloid or ctenoid or mix of both. Preopercular scales
usually cycloid, upper or rear part of preopercle may include ctenoid scales; cycloid scales may extend anterior to
vertical dark cheek bar below eye. Breast with cycloid scales reaching up to below mid-opercle or as far forward as
rear edge of preopercle. Pectoral fin base covered with cycloid scales, some may be ctenoid. Belly scales along
midline usually cycloid; ctenoid scales present posteriorly on midline in many specimens.
FIGURE 4. Diagram showing variation in ctenoid and cycloid scale arrangement on the predorsal (nape) region in specimens
of A-B, Gnatholepis anjerensis from Mauritius and Ambon respectively, C-D, G. c a u e r e n s i s , both from Aliwal Shoal, South
Africa and E-F, G. t h o m p s o n i from Tobago and Brazil respectively. Hatching indicates ctenoid scales.
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Head pores with anterior nasal pore just anterior to anterior naris, posterior nasal pore beside each posterior
naris, a pair of anterior interorbital pores, a single posterior interorbital pore, a postorbital pore, a terminal pore
over opercle and an anterior and posterior temporal pore in short separate posterior portion of the oculoscapular
canal over opercle; three preopercular pores present (Fig. 1).
Sensory papillae arranged in transverse pattern (Fig. 1); rows on opercle may be partly concealed by scales;
papillae on chin in two short longitudinal rows on either side of midline (see Larson & Buckle 2005: fig. 2B).
Coloration of fresh material. Fresh-dead specimens are illustrated in Randall and Greenfield (2001: pl.
IA–F), while good illustrations of living fish are in Senou et al. (2004: 244–245), Kuiter and Tonozuka (2001: 672,
as G. d e l t o i d e s and G. sp.1) and Myers (1991: plate 124A, 1999: plate 161D). There is a little variation in colour
pattern (Figs 5–6).
FIGURE 5. Gnatholepis anjerensis, 2 m depth, Shams Alam, south of Marsa Alam, Egypt, Red Sea. Photo by Sergey
Bogorodsky.
Living fish show characteristic small bright blue-white to pearly white spots and blotches on the side of the
head (may form oblique to horizontal rough lines on cheek and opercle), pectoral fin base and scattered along the
middle of the side of the body, mostly immediately behind the pectoral fin or less often, extending down the
anterior half of the body (Fig. 5). A characteristic pale to deep gold or orange spot is present above the pectoral fin
base, that may be surrounded by a grey to purplish grey to brownish blotch or a short brownish horizontal line, with
one or several short blue-white to pearly white short horizontal lines beneath (Fig. 6). Body colour pattern similar
to preserved specimens but blotches and short lines on body are purplish or reddish brown to orange brown in
colour, with the large lateral blotches or bars being browner than the overlying short lines and spots. The first
dorsal fin always has three to four wavy blackish to reddish-brown lines beginning on the first spine; with a
characteristic black spot on the front of the spine. The pectoral fin is finely banded or speckled with rows of dark
brown and white, giving a distinctive appearance.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
FIGURE 6. Gnatholepis anjerensis, Kosi River mouth, Kwa Zulu Natal, South Africa. Photo by Dennis King.
Coloration of preserved material. The illustrations in Randall and Greenfield (2001) show some of the
variations in colour pattern. The basic pattern is as follows.
Head and body yellowish white to pale brown (depending upon preservation) with five (sometimes indistinct)
brown saddles crossing dorsum, posterior part of each saddle merging into one of six vertical broad diffuse brown
bars or elongate to rounded blotches along mid-side of body, posteriormost bar (on caudal peduncle just before
mid-base of caudal fin) often indiscernible. Two to four irregular rows of small dark brown spots along dorsal
quarter of body, mid-side of body may be marked by series of short brown horizontal lines and spots, which may be
enlarged and merging with underlying vertical diffuse brown bars. Specimens may be heavily spotted, with most
scales on upper two-thirds of body having a brown spot.
Predorsal, snout and side of head finely speckled and spotted with brown, opercle with brown line extending
from centre of preopercular margin obliquely back to rear opercular corner. Blackish to dark brown cheek-bar from
ventral edge of eye running vertically to slightly obliquely down to end on lower preopercular edge, but not
extending onto branchiostegal membranes. Blackish to dark brown line from (usually) rear half of iris running over
top of eye, rarely meeting its counterpart in interorbital space. Upper lip crossed by dusky bar (at level with nares)
and similar dusky bar crossing centre of lip; bars on lip may be marbled, vermiculate or broken into series of
irregular spots and short streaks. Cheek with diffuse brownish horizontal bar crossing vertical cheek-bar; brownish
bar variable in shape and intensity, its pigment often surrounding the round pale spots that were white when fish
alive. Above pectoral fin base, brownish to dark brown blotch or short horizontal bar (may be diffuse to nearly
indiscernible) surrounding small pale round spot (not always discernible in preserved material); brown blotch or
bar may be partly connected to brown line or series of brown spots running from behind eye along top of
preopercle. Pectoral fin base with horizontal brown line across dorsal half, line extending onto lower part of fin and
ending in diffuse brown blotch.
First dorsal fin transparent, with three to four wavy brown lines (may break up into series of spots)
beginning at first spine; beginning of first three wavy lines marked by characteristic darker brown to blackish
spot on front of spine. Second dorsal fin similar but spots on face of fin spine much less distinct; wavy lines may
transform into many series of spots or coalesce into plain brown pigment, which may form vertical streaks on
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membrane, especially on posterior half of fin. Anal fin plain dusky to brownish, may be darker distally. Caudal
fin transparent to translucent brownish with pale brown streaks or rows of fine brown spots along fin
membranes, especially on dorsal half of fin. Pectoral fins transparent to translucent dusky, distinctively finely
speckled with darker brown, in irregular rows. Pelvic fins and frenum plain dusky to brownish, darker brown
toward midline of fused fins.
TABLE 9. Measurements of specimens of Gnatholepis anjerensis, expressed as percentage of standard length (SL) or
head length (HL); n = 60.
Distribution. Gnatholepis anjerensis is known from South Africa, Mozambique, Zanzibar, Kenya,
Madagascar, Aldabra, Seychelles, Rodrigues, Mauritius, Maldives, Chagos, Egypt, Israel, Oman, Yemen, Thailand,
Malaysia, Indonesia, Philippines, Japan, Guam, Micronesia, Marshall Islands, Vanuatu, Australia (Cocos-Keeling
Islands to One Tree Island), Solomon Islands, New Caledonia, Tonga, Fiji, Wallis Island and the Society Islands.
Ecology. Very little is reported of the natural history of this species other than Randall and Greenfield (2001),
who provided information on preferred habitat (though this data is mixed with that for other species). Gnatholepis
anjerensis is usually found in lagoonal situations over sand near coral or coral rock, rubble, Halimeda or seagrasses
at a depth range of 0.25–12 m. Coene and Ollevier (2000) found G. a n j e r e n s i s in Kenya mostly at sites with
seagrass, feeding on a general range of benthic invertebrates.
Cole (1990), in her discussion of hermaphroditism in some coral reef gobies, confirmed that Gnatholepis
anjerensis was a gonochore and did not change sex (a feature of the subfamily Gobionellinae). Her collection of 69
females and 54 males showed a sex ratio of 1.3 females per male.
Thacker’s (2004a) work on the population structure of South Pacific G. a n j e re n s i s and G. c a u e re n s i s (which
she refers to as G. s c a p u l o s t i g m a ) showed that the populations of G. a n j e r en s i s from Fiji to the Tuamotus were all
much younger (about 88,000 years) than the ages of the island groups in which they were found. Population
expansion for Gnatholepis cauerensis, on the other hand, was estimated to have occurred much earlier, about
237,000 years ago.
Neotype
(male)
Means
(males)
Maximum
(males)
Minimum
(males)
Means
(females)
Maximum
(females)
Minimum
(females)
Standard length 31 42.3 84.0 27.5 39.9 65.0 26.0
Head length in SL 30.0 29.1 31.0 26.6 29.7 33.3 27.5
Head depth in HL 66.7 71.7 83.7 63.8 70.9 82.1 62.0
Head width in HL 67.7 67.9 82.2 53.7 69.4 79.5 62.1
Body depth at anus in SL 22.3 23.0 26.3 20.6 23.7 26.6 19.6
Body depth at D1 origin in SL 22.6 23.4 25.8 14.3 23.3 27.7 14.6
Caud. ped. length in SL 14.8 15.7 18.2 12.9 16.4 18.6 14.7
Caud. ped. depth in SL 11.3 11.9 13.8 10.9 11.6 13.0 10.4
Snout length in HL 34.4 34.9 43.4 22.4 34.6 44.4 29.2
Eye width in HL 30.1 26.8 29.8 22.7 27.7 30.5 24.0
Jaw length in HL 34.4 36.9 41.5 32.7 34.7 38.1 32.0
Interorbital in HL 8.6 8.4 11.3 6.1 7.2 9.4 4.7
Preorbital width in HL 21.5 22.4 27.8 18.2 22.2 26.5 18.4
Pectoral fin in SL 27.1 25.9 28.7 23.6 26.1 28.3 23.5
Pelvic fin in SL 27.1 28.2 31.7 22.1 26.3 29.8 22.4
Caudal fin in SL 29.4 32.0 36.1 25.1 29.5 32.5 25.8
Adpressed D1 in SL 23.5 25.1 28.5 18.9 24.6 27.9 21.2
3
rd
D1 fin spine in SL 16.1 17.9 19.3 16.0 17.2 19.6 15.4
4
th
D1 fin spine in SL 15.2 18.1 19.8 16.0 17.2 19.6 15.0
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Comparisons. When alive, this species is most easily distinguished from its congeners by having the first
dorsal fin rather square to rounded in shape (the fifth D1 spine is nearly as long as the third to fourth spines), in
having the first spine banded by three to four black spots or short streaks; the pectoral fin is covered with fine white
and dark speckles, a characteristic small round gold spot is above the pectoral fin base (often with a short dark line
or elongate blotch under it) and the nape and body is covered with irregular rows of small blackish to reddish spots,
with five to six diffuse dusky blotches along the lower mid-side of the body. When preserved, G. a n j e r e n s i s can be
distinguished by its ctenoid predorsal scales, slightly stocky body and square first dorsal fin shape; most of the
colour pattern remains visible but for the small round gold spot (which may be hard to discern) above the pectoral
fin base.
Remarks. Gobius anjerensis was based on a Kuhl and van Hasselt figure of a 48 mm specimen, and is the type
species of the genus Gnatholepis. A neotype (BPBM 26651) was designated by Randall and Greenfield (2001: 3,
fig. 2). Randall and Greenfield (2001) picked a fairly small (31 mm SL; the species reaches 84 mm SL) and slightly
fragile specimen as neotype of Gobius anjerensis (from northern Sulawesi, some distance from the type locality of
Anjer in Java, but forgivable in that Anjer was destroyed in the 1883 eruption of Krakatau). Some of Randall and
Greenfield’s counts do not agree with ours. For example the lateral scale count for the neotype is given as 30, but
we counted 26 scale rows on this specimen and their Figure 2 of the neotype also shows 26 scales. Although
Randall and Greenfield (2001) say that they counted lateral scales “from the upper edge of the gill opening”, they
also say that scales are crowded behind the gill opening and that they counted all the small scales here. It appears
that they were counting lateral rows beginning right behind the gill opening and thus their counts would be several
rows greater than ours.
Gobius capistratus Peters, 1855, was described from Ibo, Mozambique, with two syntypes (ZMB 2103).
Randall and Greenfield (2001) synonymised this with G. a n j e re n s i s but gave no reason for doing so and apparently
did not examine type material. Peters’ specimens were described and figured in Sauvage (1891: 356, pl. 38, fig. 5)
and a partial translation of Peters’ description was provided in Günther (1861: 36). Klunzinger (1871) considered
that G. o p h t h a l m o t a e n i a was a synonym of G. c a p i s t r a t u s . Jeff Leis photographed the syntypes at ZMB and the fine
light and dark speckling on the pectoral fins is visible, as are the blue-white spots on the lower part of the body and
side of the head (in both specimens) (Fig. 7).
Gobius deltoides Seale, 1901, was described from 18 syntypes from Guam (ANSP 84134, BPBM 267, USNM
109399). ANSP 84134 (formerly BPBM 267) includes a 14 mm characoid.
Some specimens identified by Randall and Greenfield as G. a n j e r e n s i s (AMS I.20757-072, Raine Island;
AMS I.27156-035, Middleton Reef; AMS IB.4004, Heron Island), were subsequently identified by us as G.
cauerensis (n.b. one specimen in AMS I.20757-072 was G. a nj e re n s i s , the other 10 were G. c a u ere n si s ). These
may be some of the anomalous specimens that they mentioned as having distinct longitudinal lines on the body,
a colour pattern not usually found in G. a n j e re n si s (Randall & Greenfield 2001: 4–5). Their definition of the
species G. a n j ere ns i s should be taken from the description of the neotype only, as their view of this species is
confused.
Thacker’s (2004b) definitions of G. a n j e r e n s i s and G. k n i g h t i do not agree with ours. From her discussion
(Thacker 2004b: 578–580, fig. 3), it seems that she based her decision as to what name with which to label each
taxon, on whether or not the fish had a dark blotch or spot, with or without a pale centre, above the pectoral fin base
(these she called G. s c a p u l o s t i g m a ), a dark line or dash above the pectoral fin base (these she called G. k n i g h t i ), or
had no distinct spot at all above the pectoral fin base (these she called G. a n j e re n s i s ). Specimens identified as G.
anjerensis from AMS by Thacker, were re-examined by AMS fish section staff on our behalf and were found to
include Gnatholepis gymnocara (specimens from Yeppoon, Sabina Point and Townshend Island, all in
Queensland). Two lots from the Northern Territory (East Arm; Gunn Point) were found to be G. a rg u s , not G.
anjerensis, as pointed out by Larson and Buckle (2005).
Kuiter and Tonozuka (2001: 671, figs C, D) show specimens from Gilimanuk and Lipah, Bali, Indonesia,
which appear to be oddly-marked specimens of G. a n j e re n s i s . They show the black spots along the leading edge of
the first dorsal spine but the fin is patterned with rows of oval spots and the body is very darkly mottled. Otherwise
they closely resemble G. y o s h i n o i .
Hayashi and Shiratori’s Figure 176 is of G. a n j e re n s i s (2003: 96), while Figure 175 is of G. y o s h i n o i (see
Remarks for that species).
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FIGURE 7. Gnatholepis anjerensis, syntypes of Gobius capistratus, ZMB 2103. Photo by Jeff Leis.
Gnatholepis argus Larson and Buckle, 2005
(Fig. 8; Tables 5–8)
Gnatholepis gymnocara Randall and Greenfield, 2001: 14 (in part).
Gnatholepis sp.—Randall and Greenfield 2001: 16, pl. IIH (Northern Territory); Thacker 2004b: figs 1–2.
Gnatholepis argus Larson and Buckle, 2005: 68–72 (N side of Turtle Reef off Field Island, Northern Territory).
Diagnosis. (From Larson and Buckle 2005). A small Gnatholepis (up to 33 mm SL) lacking scales on side of
head, predorsal midline and pectoral fin base, no distinct flap on end of lower lip (low fold may be present), and
no distinct canine or enlarged teeth; sexually dimorphic in adult size and colour, with females averaging larger
than males; sexually dichromatic: males with row of six dusky to blackish spots or blotches along mid-side of
body, posteriormost 3–4 blackish spots darker and larger than anterior spots and surrounded by small iridescent
blue spots in life, small dense black spots scattered on unpaired fins; females with broken grey line along mid-
side of body and dusky to blackish spots or blotches indistinct and posterior blotches not darker than anterior
ones, no small iridescent blue spots on body; second dorsal fin rays modally I,10; anal fin rays modally I,11;
pectoral rays 15–18; lateral scales 24–28; predorsal scales absent from nape midline, scales on side of head may
reach from just over opercle to nearly behind eyes (Fig. 8).
Remarks. As pointed out by Larson and Buckle (2005), Randall and Greenfield (2001) included two
paratype specimens of G. g y m no c a ra in their description (AMS I.23930-011). As no other specimens of G.
gymnocara have so far been found in the Territory, these paratypes were examined and were found to be female
G. arg u s.
The range of this species includes Western Australia, from Cape Talbot south to Enderby Island, Dampier
Archipelago; also Daru, Papua New Guinea (specimens at Western Australian Museum, Perth).
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
FIGURE 8. Gnatholepis argus, fresh dead colour, male, AMS I.24677-003, Bullocky Point reef, Darwin, Northern Territory,
Australia. Photo by Doug Hoese.
Gnatholepis caudimaculata new species
(Figs 9–11, 12A; Tables 5–8, 10)
Gnatholepis anjerensis—Randall 1995: 336 (Oman); Debelius 1998: 180 (Na’ama Bay, Sinai).
Diagnosis. A moderate-sized Gnatholepis (up to 42.5 mm SL) with cycloid scales on head and pectoral fin base,
predorsal midline usually naked, distinct flap present at end of lower lip; when alive, body whitish with finely
speckled dorsum, characteristic vertical oval blackish to dark grey blotch across caudal fin base, two broad ladder-
like orange to reddish stripes along body and oblique black curved or comma-shaped mark on top of eye; second
dorsal and anal fin rays nearly always I,11; pectoral rays 15–17, modally 16; lateral scales 25–28, modally 26; 7–10
predorsal scales (usually 8).
Material examined. HOLOTYPE—USNM 327425, 45 mm SL male, reef near road at Marsa Muqabila,
north-west coast Israel, Gulf of Aqaba, V.G. Springer and party, 17 July 1969. PARATYPES—BPBM 27442,
4(18.5–28), Towartit Reef, Sudan, Red Sea, J.E. Randall, 13 January 1980; PMR VP.2219, 2(26.5–30.5),
Sanganeb, Sudan, Red Sea, S. Bogorodsky, 11 October 2009; PMR VP.2220, 1(28.5), Shaab Suadi, Sudan, Red
Sea, S. Bogorodsky, 12 October 2009; USNM 313779, 1(28.5), bay on NW end of Giftun Al Kebir Island,
Hurghada, Strait of Jubal, Egypt, H.A. Fehlmann and party, 3 January 1965; BPBM 13401, 1(34.5), Dahab, Gulf of
Aqaba, Egypt, J.E. Randall, 14 June 1972; USNM 313780, 21(19.5–29), S shore Difnein Island, Ethiopia, V.G.
Springer and party, 15 August 1969; USNM 399290, 11(16–36), reef near road at Marsa Muqabila, north–west
coast Israel, Gulf of Aqaba, V.G. Springer and party, 17 July 1969; BPBM 18231, 11(26–37.5), 1 km N of Coral
Island, Gulf of Aqaba, J.E. Randall and O. Gon, 23 September 1974; ex-USNM 327464, 3(27–31.5), Zubair Island,
Yemen, E. Clark, 30 September 1967; BPBM 33350, 2(30–36.5), SE side Jana Island, Saudi Arabia, Persian Gulf,
J.E. Randall and party, 13 September 1985; BPBM 33373, 1(42.5), S side Jana Island, Saudi Arabia, Persian Gulf,
J.E. Randall and party, 13 September 1985; BPBM 33374, 2(31–36.5), Jana Island, Saudi Arabia, Persian Gulf, J.E.
Randall and party, 13 September 1985; BPBM 30457, 3(33.5–35), N side Jana Island, Saudi Arabia, Persian Gulf,
J.E. Randall and party, 15 June 1984.
Non-type specimens examined. BPBM 13423, 2, El Hamira, Gulf of Aqaba, Egypt; BPBM 18358, 1, Ras
Mohammed, S end of Sinai Peninsula, Egypt.
Description. Based on 37 specimens, 26–42.5 mm SL. An asterisk indicates the counts of the holotype (Fig.
9).
First dorsal VI*; second dorsal I,10–I,11* (mean I,10.9); anal I,10–12 (mean I,10.9, I,11 in holotype), pectoral
rays 15*–17 (mean 15.8), segmented caudal rays always 17*; caudal ray pattern always 9/8*; branched caudal rays
7/6 to 7/7* (usually 7/6); lateral scale count 25–28 (mean 26.3, 26 in holotype); TRB 8–11 (mean 10.0*); predorsal
scales 6–10 (mean 8.1, 8 in holotype); circumpeduncular scales always 12*. Gill rakers on outer face of first arch 1
+ 3–4 (in 4, usually 1+4).
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FIGURE 9. Holotype of Gnatholepis caudimaculata n. sp., USNM 327425, 36 mm SL male, Marsa Muqabila, Israel, Gulf of
Aqaba. White object in mouth is paper label; pinkish cast to first dorsal fin due to small pebble holding specimen upright.
Body compressed, width at anus 9.0–15.9% (mean 13.1%) of SL. Body rather stocky in appearance, body
depth at anus 13.4–24.6% (mean 22.0%) of SL, body depth at first dorsal fin origin 21.0–25.7% (mean 23.3%) of
SL. Head compressed, slightly broader ventrally, slightly deeper than wide, HL 24.6–30.7% (mean 28.4%) of SL;
head depth at posterior preopercular margin 62.8–82.4% (mean 69.8%) of HL; head width at posterior preopercular
margin 55.8–88.4% (mean 67.6%) of HL; head profile bluntly pointed to somewhat rounded; nape profile straight
to slightly curved. Mouth nearly terminal, fleshy snout tip may slightly overhang tip of jaws, slightly oblique; jaws
reaching to anterior half of eye or just below anterior margin of eye in large specimens; upper jaw length
32.1–39.8% (mean 35.4%) of HL. Upper lip smooth, narrower than lower, lower lip papillose close to teeth, with
ridge of small papillae along side of lip ending in twist or fold posteriorly, forming triangular flap, lip interrupted at
chin. Eye moderate, dorsolateral, 25.0–32.6% (mean 28.6%) of HL; preorbital width 14.6–21.7% (mean 19.0%) of
HL. Snout usually bluntly pointed, 26.3–38.2% (mean 32.1%) of HL; posterior naris rounded, close to mid-level
margin of eye; anterior naris in short tube, about level with ventral half to middle of eye. Interorbital narrow,
5.8–12.2% (mean 7.8%) of HL. Caudal peduncle compressed, length 14.4–19.5% (mean 17.0%) of SL; caudal
peduncle depth 9.8–11.9% (mean 11.0%) of SL.
First dorsal fin rounded to almost triangular, with no spines greatly elongate; second to fourth spine usually
longest; when adpressed, spine tips reaching to first to third element of second dorsal fin in males, fin may fall short
in females. Second dorsal spine length 13.4–19.1% (mean 17.1%) of SL; third dorsal spine length 13.4–21.8%
(mean 17.8%) of SL; fourth dorsal spine length 13.1–19.7% (mean 17.3%) of SL. Second dorsal fin nearly as tall as
first dorsal fin anteriorly, fin pointed posteriorly. Anal fin lower than second dorsal fin, anteriormost rays shorter
than posterior rays; fin pointed posteriorly. Second dorsal and anal fin rays, when adpressed, reaching caudal fin in
males, usually falling just short of fin ray bases in females. Pectoral fin oval to somewhat pointed, central rays
longest, 21.4–28.8% (mean 24.8%) of SL; fin reaching back to just above anal fin spine. Pelvic fins fused, frenum
with distinctive finely fimbriate margin, fins round to oval, reaching to anal fin spine, 23.6–30.7% (mean 26.7%) in
SL. Caudal fin moderate, rounded posteriorly, 25.2–33.0% (mean 29.8%) of SL.
Gill opening restricted, extending anteriorly to lower edge of pectoral base. One or two slender gill rakers on
outer face of first arch, closest to angle of arch, remainder of rakers very short and stubby; clumps of tiny pointed
papillae present along inner face of first arch; outer rakers on second gill arch mostly replaced by several series of
fine papillae; outer rakers on remaining arches low, broad-based. Inner face of upper limb of first gill arch, and to
lesser extent, upper limbs of other arches, covered with low dense fleshy papillae which may form clumps or
groups; dorsal portion of arch may have short fleshy protuberances ending in one or several papillae. About one-
third of first gill arch bound by membrane to inner face of opercle. Tongue tip bilobed.
Teeth in upper jaw in two to three rows across front and one row at side of jaw, outermost row teeth largest,
curved and pointed, largest teeth at front of jaw on either side of symphysis; innermost row teeth small, sharp and
evenly sized. Teeth in lower jaw in two to three rows, arranged similarly to upper jaw but outer row teeth smaller in
females; posteriormost one or two outer row teeth enlarged or recurved in males.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
FIGURE 10. Gnatholepis caudimaculata n. sp., 9 m depth, north of Marsa Alam, Egypt, Red Sea. Photo by Sergey
Bogorodsky.
Predorsal scales cycloid, of variable extent, from just over opercle to (rarely) close up to behind eyes; nape
midline usually naked, with scales present on either side of nape. Opercle covered with scales, cycloid or ctenoid or
mix of both. Preopercular scales cycloid; occasionally few scales present anterior to vertical dark cheek-bar below
eye. Breast with cycloid scales usually reaching up to below mid-opercle. Pectoral fin base with cycloid scales.
Belly scales along midline usually cycloid; few ctenoid scales may be present anterior to anus.
Sensory papillae and head pores as in G. a n j e re n s i s .
Coloration of fresh material. Photographs of living fish (as G. a n j e re n s i s ) appear in Randall (1995) and
Debelius (1998). The following description is mostly based on photographs of living fish by Sergey Bogorodsky
and Ole Brett (Figs 10–11).
Living fish colour pattern very similar to preserved pattern (see below). Head and body white to slightly
greyish white with two broad ladder-like orange to brownish stripes formed by two rows of small rounded spots
along body, mid-lateral stripe usually most pronounced and upper stripe (commencing at upper corner of opercle)
less distinct to nearly absent; six diffuse to indistinct small brownish blotches interspersed with white patches
evenly spaced along lower stripe; most distinct marking a vertically-oriented black or blackish oval to rectangular
blotch on mid-base of caudal fin at hypural crease.
Dorsum lightly speckled with small light to dark brown spots and a few small orange spots; may be up to eight
evenly spaced patches of bluish white placed above dorsalmost ladder-like orange stripes (or rows of spots). Snout
lightly mottled with pearly pale bluish white with some pale diffuse brown mottling near nares. Opercle crossed by
bright orange oblique stripe, ending near upper rear corner, may be some light brown mottling around stripe; cheek
sometimes with brown mottling around vertical to slightly oblique dark orange to red-brown or dark brown narrow
bar or line running from eye down to lower preopercular edge, or cheek only with vertical bar from eye and a short
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horizontal brownish mark extending anteriorly from rear edge of preopercle; characteristic very dark brown to
black oblique to comma-shaped mark across top of eye. Brown-edged bright orange and brown line from behind
eye running atop preopercle, opercle and across pectoral fin base (anteriormost part of line usually brown); forming
lower edge of diffuse dark brown broadly-forked blotch above fin base and yellow to pale gold spot present in
centre of blotch. Pectoral fin base whitish speckled with brown, and brown-edged bright orange bar crossing just
above centre of fin base. Upper lip crossed by dark brown line at level of nares and smaller brown or diffuse brown
marks may be present at midline of lip. Lower lip, chin and breast white. Iris silvery to pale golden, with brown line
(from cheek-bar) ventral to pupil and red-brown to black line above pupil that curves around eyeball in comma-
shape posteriorly.
FIGURE 11. Gnatholepis caudimaculata n. sp., Sharm, Sinai, Egypt. Photo by Ole Brett.
First dorsal fin translucent to transparent, crossed by narrow irregular brownish lines or series of small
brown spots. Second dorsal fin very similar but with series of elongate to oval darker brown spots toward
posterior half. Anal fin translucent whitish with rows of irregular small brown spots; may become more elongate
posteriorly. Caudal fin translucent white to transparent, crossed by irregular rows of small brown spots and short
streaks and vertically-oriented black or blackish oval to rectangular blotch on mid-base of fin. Pectoral fins
transparent, lightly speckled with bright white on basal half and with small diffuse brownish blotch on bases of
rays near middle of fin (continuation of horizontal orange bar on fin base). Pelvic fins transparent to translucent
whitish.
Coloration of preserved material. Head and body yellowish white to whitish, with two broad ladder-like
light brown stripes (about one scale wide) along body, first stripe beginning at upper rear corner of opercle and
ending on upper surface of caudal peduncle just before fin base, second stripe beginning behind pectoral fin base
and ending on lower half of caudal fin base; stripes with slightly darker upper and lower margins; similar stripe
along dorsum from origin of first dorsal fin to middle of second dorsal fin (may be replaced by irregular brown
speckling or series of about eight diffuse short faint brownish saddles over dorsum); vertically-oriented brownish
oval to rectangular blotch on mid-base of caudal fin (at hypural crease).
Predorsal lightly speckled with fine melanophores or scattered brown spots; snout generally pale with
brown mottling near nares; opercle mottled with light brown, some specimens with dusky-edged pale oblique
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
stripe crossing opercle, ending near upper rear corner; cheek with brown mottling (which may curve around a
few unpigmented spots) and vertical to slightly oblique dark brown narrow line running from eye down to
lower preopercular edge; distinctive dark brown to blackish oblique to comma-shaped mark across top of eye,
beginning above mid-pupil and ending past rear edge of pupil (nearly to rear edge of eye in some specimens)
(Fig. 12A). Variably intense brownish line running from behind eye atop preopercle and opercle, across
pectoral fin base and forming lower edge of pale to dark brown broadly-forked blotch above fin base
(unpigmented area usually present in centre of ‘fork’). Pectoral fin base pale to brown-speckled, with darker
brown bar crossing just above centre of fin base. Upper lip crossed by brown line or blotch at level of nares
and smaller brown spot may be present at midline of lip. Lower lip, chin and breast whitish or lightly dusted
with melanophores.
First dorsal fin translucent whitish, crossed by five to six narrow irregular brownish lines, which may be
partly broken-up by series of dark spots posteriorly; fin margin faintly dusky. Second dorsal fin very similar but
wavy lines mostly broken up into series of elongate to oval dark spots. Dorsal fins more strongly marked in
males. Anal fin transparent to translucent with broad dusky margin. Caudal fin translucent whitish to pale
yellowish brown, crossed by about eight irregular rows of small brown spots and short streaks; in many
specimens most of these markings ‘blurred’ and diffuse except for on upper and lower parts of fin; males with
most pronounced spotting. Pectoral fins transparent to lightly speckled with brownish pigment and brownish
blotch on bases of rays near middle of fin (continuation of horizontal brown bar on fin base). Pelvic fins
transparent to brownish; frenum pale.
Distribution. Restricted to the Red Sea (Ethiopia, Israel), Gulf of Aqaba and Persian Gulf (Saudi Arabia). The
species has been reported from Oman and Egypt, although identified as G. a n j e re n s i s (Randall 1995; Debelius
1998).
Ecology. Very little information available; depth range of material examined was 0–17 m. Photographs of
living fish show them over white to blackish sand. This species has been shown to exhibit red fluorescence on the
dorsal half of the eye by Michiels et al. (2008: fig. 5; fish identified as G. a n j e re n s i s ). They identified 20 species of
red-fluorescing gobiids (of the genera Bryaninops, Ctenogobiops, Eviota, Fusigobius, Gnatholepis, Istigobius,
Pleurosicya and Trimma). This has been hypothesised to be part of a “private communication system” as the
fluorescence occurs in features that are used for intra-specific signals, especially the eyes. As Gnatholepis species
have distinctive and slightly different black markings across the dorsal surface of the eyeball, this red fluorescence
and the pattern on the eye may be of considerable significance to the fish.
Comparisons. This species is very like G. c a u e re n s i s , and replaces that species in the Red Sea. It is
distinguished by possession of a vertical black to dark brown blotch on caudal fin base, the ladder-like rows of
small spots forming two stripes along the body and the obliquely placed comma-shaped lines atop each eye. It has
a slightly lower number of predorsal scales compared to G. c a u e re n s i s : 7–9, modally 8, compared to 7–12, modally
9 (comparing data from 27 G. c a u d i m a c u l a t a specimens to 113 G. c a u e re n s i s ).
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FIGURE 12. A, Sketch of dorsal view of head of Gnatholepis caudimaculata n. sp., showing characteristic oblique mark on
top of eye; B, dorsal view of head of Gnatholepis anjerensis; C, dorsal view of head of Gnatholepis cauerensis; D, dorsal view
of head of Gnatholepis knighti; E, dorsal view of head of Gnatholepis ophthalmotaenia; F, dorsal view of head of Gnatholepis
thompsoni.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
TABLE 10. Measurements of specimens of Gnatholepis caudimaculata n. sp., expressed as percentage of standard
length (SL) or head length (HL); n = 33.
Gnatholepis cauerensis (Bleeker, 1853)
(Figs 4C–D, 12C, 13–18; Tables 5–8, 11)
Gobius cauerensis Bleeker, 1853: 269 (Cauer, Sumatra, Indonesia).
?Gobius knighti—Aoyagi 1957: 228 (Japanese Archipelago).
Acentrogobius cauerensis—Palmer 1970: 227–228 (Betio, Tarawa); Goren 1979: 18 (Israel, Egypt, Eritrea, Ethiopia; Red Sea).
Gnatholepis cauerensis—Hoese and Winterbottom 1979: 4 (South Africa); Goren 1981: 98 (New Caledonia); (in part) Maugé
1986: 369 (Transkei northwards); Allen and Smith-Vaniz 1994: 16 (Cocos (Keeling)); Myers 1999: 255, plate 161E
(Micronesia); Randall 1999: 26 (Pitcairn Island); Larson in Randall and Lim 2000: 637 (South China Sea); Randall and
Earle 2000: 19 (Marquesas); Randall and Greenfield 2001: 6; Hutchins 2001: 43 (Western Australia); Larson and Murdy
2001: 3601; Kuiter and Tonozuka 2001: 673, figs A–C (Bali); Myers and Donaldson 2003: 638 (Guam); Allen and Adrim
2003: 58 (Papua to Sumatra); Galzin et al. 2002: 232 (Rapa); Manilo and Bogorodsky 2003: S118; Randall et al. 2004: 27
(Tonga); Lobel and Lobel 2004: 76 (Wake Atoll); Heemstra et al. 2004: 3329 (Rodrigues); Anderson 2005: 97 (Maldives);
Hoese and Larson 2006: 1654; Williams et al. 2006: 259 (Wallis Islands); Fricke et al. 2009: 101.
Gnatholepis anjerensis—Dor 1984: 237 (Red Sea); Randall et al. 1985: 70 (Johnston Island); Randall et al. 1990b: 33, fig. 43 (Rapa).
Gnatholepis scapulostigma Herre, 1953: 193 (Engebi Island, Eniwetok [Eniwetak] Atoll, Marshall Islands).—Yoshino in
Masuda et al. 1984: 252 (Ryukyu and Ogasawara Islands); Randall and Randall 1987: 309 (Enewatak Atoll); Randall et al.
1990a: 402; Myers 1991: 238, 240, pl. 124B); Randall and Goren 1993: 13 (Maldives); Kuiter 1993: 347 (New South
Wales); Allen 1997: 216; Kulbicki and Williams 1997: 22 (Ouvéa Atoll, New Caledonia); Fricke 1999: 513 (Réunion);
Laboute and Grandperrin 2000: 411 (New Caledonia); Larson and Murdy 2001: 3601; Kuiter and Tonozuka 2001: 673,
figs A–C (Maumere, Flores); Nakabo 2002: 1212; Hayashi and Shiratori 2003: 97.
Gnatholepis inconsequens Whitley, 1958: 44 (Heron Island, Capricorn Group, Queensland).—Allen and Swainston 1988: 134
(Rottnest Island); Gill and Reader 1992: 223 (Middleton and Elizabeth Reefs); Francis 1993:167 (Lord Howe Island);
Larson and Murdy 2001: 3601.
Gnatholepis cauerensis australis Randall and Greenfield, 2001: 7, pl. IIB (Haurei Bay, Rapa, Tubuai Islands).
Holotype
(male)
Means
(males)
Maximum
(males)
Minimum
(males)
Means
(females)
Maximum
(females)
Minimum
(females)
Standard length 41.0 32.8 42.5 26.5 30.0 36.5 26.0
Head length in SL 26.8 28.2 30.7 26.4 28.7 30.0 24.6
Head depth in HL 68.2 69.1 82.4 61.4 70.6 78.7 66.7
Head width in HL 70.0 67.3 88.4 55.8 67.9 76.0 59.3
Body depth at anus in SL 22.2 21.5 24.2 13.4 22.5 24.6 19.0
Body depth at D1 origin in SL 22.2 22.8 25.5 19.5 23.9 25.7 21.0
Caud. ped. length in SL 17.6 17.2 19.5 15.1 16.8 19.1 14.4
Caud. ped. depth in SL 10.7 11.2 11.9 10.4 10.8 11.6 9.8
Snout length in HL 35.5 32.5 38.2 26.3 31.6 35.0 27.7
Eye width in HL 27.3 28.1 30.8 25.7 29.2 32.6 25.0
Jaw length in HL 38.2 35.6 39.8 33.3 35.3 39.5 32.1
Interorbital in HL 9.1 7.8 10.1 5.8 7.9 12.2 6.5
Preorbital width in HL 20.9 18.9 21.6 15.1 19.1 21.7 14.6
Pectoral fin in SL 22.4 24.9 28.8 22.4 24.6 27.2 21.4
Pelvic fin in SL 27.8 27.3 30.7 24.3 25.9 28.2 23.6
Caudal fin in SL 30.5 30.6 33.0 28.2 28.8 30.7 25.2
Adpressed D1 in SL 24.6 25.1 27.7 21.8 23.7 28.6 18.4
3
rd
D1 fin spine in SL 18.3 18.1 20.0 16.1 17.5 21.8 13.4
4
th
D1 fin spine in SL 17.6 18.0 19.7 16.9 16.3 18.2 13.1
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Gnatholepis cauerensis hawaiiensis Randall and Greenfield, 2001: 10, pl. IIC, D (Pukukea, Oahu, Hawaiian
Islands).—Greenfield and Randall 2004: 526–528 (Hawaiian Islands).
Gnatholepis sp.—Wass 1984: 28 (Samoa); Myers 1991:240, pl. 124C).
Coryphopterus—Lobel 2003: 113 (Johnston Atoll) [pasted-in errata sheet on inside back cover says Gnatholepis anjerensis].
Gnatholepis cauerensis cauerensis—Senou et al. 2004: 242; Lobel and Lobel 2004: 76 (Wake Atoll); Motomura et al. 2010:
213, fig. 525 (Yaku–shima Island).
Gnatholepis australis—Randall 2009: 180, fig. 3.
Gnatholepis hawaiiensis—Randall 2009: 181.
Diagnosis. A moderate-sized Gnatholepis (up to 56 mm SL) with wide range in live colour pattern; ctenoid body
scales usually reaching rear corner of opercle, occasionally ctenoid scales reaching to above rear preopercular
margin; nape midline scales always cycloid, distinct flap present at end of lower lip; when alive, colour variable:
body translucent white with five to seven rows of small variably coloured (dull orange, brown, red-brown to
blackish) spots and/or thin lines and about six indistinct diffuse orange-brown to purple-brown blotches along mid-
side; narrow vertical black mark on top of each eye, marks may join each other across interorbital space, and
narrow black vertical line crossing cheek below eye; small dark mark of variable shape (often W-shaped) above
pectoral fin base, with small yellow spot in centre; second dorsal and anal fin rays usually I,11; pectoral rays
12–18, modally 16; lateral scales 24–30, modally 27; 7–12 predorsal scales (modally 9).
FIGURE 13. Gnatholepis cauerensis, Scott Reef, Western Australia. Photo by Gerry Allen.
Material examined. SOUTH AFRICA: SAIAB 62111, 4(30–43), “Tiger Cove”, Aliwal Shoal, Kwa-Zulu
Natal, 7 March 2000; SAIAB 60280, 1(46), Aliwal Shoal, Kwa-Zulu Natal, 23 April 1999; SAIAB 70985, 1(45),
Manta Point, Aliwal Shoal, Kwa-Zulu Natal, 21 May 2002; SAIAB 73710, 7(16–40.5), Manta Point, Aliwal Shoal,
Kwa-Zulu Natal, P. Heemstra, S. Warren, J. Stapley, 22 May 2002; SAIAB 64645, 3(39–45), Sodwana Bay, Kwa-
Zulu Natal, 13 May 2001; SAIAB 69344, 3(27–40), Three-mile Reef, Sodwana Bay, Kwa-Zulu Natal, 16 May
2002; SAIAB 56561, 2(43–49), deep sponge reef, Sodwana Bay, Kwa-Zulu Natal, 12 August 1997; SAIAB 58554,
3(26.5–36.5), “The Hospital”, Aliwal Shoal, Kwa-Zulu Natal, 14 June 1998; NTM S.14694-001, 1(32), “The
Hospital” reef, Aliwal Shoal, Kwa-Zulu Natal, P. Heemstra and party, 8 February 1998. RODRIGUES: SAIAB
70332, 1(30), Yacht Casse, P. Heemstra and party, 21 October 2001. DJIBOUTI: MNHN 1977-628, 3(22–38), W
of Maskali Island, Gulf of Tadjourah, L. Maugé and J. Randall, 21 May 1977. THAILAND: BPBM 22804, 1(305),
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Ko Ming, Similan Islands, 14 February 1979. INDONESIA: RMNH 4523, holotype of Gobius cauerensis, 1(31.5),
Kauer, Sumatra, P. Bleeker, 1852-54; NSMT P.70196, 3(13–35), off NW coast Air Island, Lombok, K. Matsuura, 3
February 1994; NSMT P.70407, 2(25–29.5), Gerupuk, Lombok, K. Matsuura, 7 February 1994; USNM 210157,
45(13–33), E shore Piru Bay, Tandjung Liang, Ceram, V.G. Springer and M. Gomon, 10 January 1973. TAIWAN:
ASIZP P.56809, 3(38–39), Tauzuwan, S.C. Lee, 25 May 1987; USNM 327456, 22(26.5–52), SW shore just off
Chuan-fan-Shi, V.G. Springer and party, 30 April 1968. JAPAN: AMS I.23492-015, 1(34.5), tidepools below
airport, Miyake-jima, D.F. Hoese and Y. Yogo, 14 May 1980; AMS I.27367-007, 1(27), Amitori Bay, Iriomote-
jima, 16 February 1976; NSMT-P49186, 2(27–33), Ishigaki-jima, Yaeyama Group, Ryukyu Islands, 10 December
1995. PHILIPPINES: AMS I.21907-003, 4(16–34), S of Anilao, Batangas Province, C. Ferraris and E. Murdy, 22
April 1980; AMS I.40155-059, 1(36), S coast Ambulong Island, MIN team, 1 June 2000; AMS I.40161-079,
3(26.5–31), Mindoro Island, 3 June 2000; USNM 298581, 6(38–47), Y’ami Island, Batanes, 26 April 1987.
AUSTRALIA: AMS I.28993-009, 1, W side near Wreck Point, North Keeling Island, Cocos-Keeling Islands;
AMS I.19641-022, 2(37.5–40), off Tantabiddi Creek, Western Australia, G.R. Allen, 27 June 1975; AMS I.33749-
122, 12(19–33), E side northern lagoon, Boot Reef, Coral Sea, FNQ team, 28 January 1993; NTM S.13618-020,
8(23–34), E side northern lagoon, Boot Reef, Coral Sea, FNQ team, 28 January 1993; AMS I.20774-107,
7(24.5–40), N tip of un-named reef, Cape Melville, Queensland, AMS-AIMS Survey, 9 February 1979; AMS
I.20757-072, 10(21–36), W end of Raine Island, Queensland, AMS I.22635-005, 1(35.5), Escape Reef North,
November 1981; AMS-AIMS Survey, 13 February 1979; AMS I.17445-049, 1(42.5), C transect, One Tree Island,
F. Talbot and party, 19 September 1968; USNM 308212, 15(33–39), reef flat on SE side One Tree Island,
Queensland, V.G. Springer and party, 27 November 1966; AMS IB.3916, holotype of Gnatholepis inconsequens,
1(32.5), Heron Island, Queensland, R. Slack-Smith, 1957; AMS IB.4004, 2(37–42.5), Heron Island, Queensland,
R. Slack-Smith, 1957; AMS I.30310-034, 1(26), SW side North Solitary Island, New South Wales, 5 May 1977;
AMS I.18659-001, 1(36), Seal Rocks, New South Wales, 10 May 1985; AMS I.19500-003, 1(31.5), Parsley Bay,
Sydney Harbour, New South Wales, R. Kuiter, 2 February 1976; AMS I.27156-035, 7(33–53), lagoon, Elizabeth
Reef, Tasman Sea, S. Reader, A. Gill, D. Ledbitter, 14 December 1987; AMS I.27142-030, 6(33–55), lagoon,
Middleton Reef, Tasman Sea, AMS party, 7 December 1987; AMS I.27138-055, 11(21–44), mid-back lagoon,
Middleton Reef, Tasman Sea, S. Reader, A. Gill, D. Ledbitter, M Cordell, 5 December 1987. NEW GUINEA:
NTM S.13661-047, 5(23.5–40), outer barrier reef S of Rasch Passage, Madang, H.K. Larson and M. Jebb, 5
October 1992; NTM S.13691-016, 1(30), S end Tab Island, near Christensen Research Institute, Madang, H.K.
Larson and M. Jebb, 25 October 1992; NTM S.13685-044, 3(20–40), reef N of Tab Island, near Christensen
Research Institute, Madang, H.K. Larson and M. Jebb, 21 October 1992. MICRONESIA: USNM 224767,
4(27–30), N coast Ponape, Caroline Islands, 4 September 1980. SOLOMON ISLANDS: NTM S.12714-008, 1(37),
Munda Lagoon reef, S. Blaber, May 1989. FIJI: USNM 243062, 6(25–33.5), Yanutha Islet, Ona Llau, Lau Group
(Eastern Archipelago), 29 April 1982; BPBM 14591, 2(27.5–30.5), outside Mbengga barrier reef, Frigate Passage,
Viti Levu,11 March 1973. TONGA: USNM 340186, 1(25), Ofolonga Island, Ha’apai Group, 12 November 1993.
NEW CALEDONIA: MNHN 1980-744, 1(36), coral reef, P. Laboute and Menou, January 1979. VANUATU: AMS
I.37905-034, 1(28), Siviri village, Efate Island, J. Williams and party, 4 May 1997. KIRIBATI: USNM 116115,
1(32.5), Hull Island Channel, Phoenix Islands, L.P. Schultz, 1939. MARSHALL ISLANDS: USNM 34429,
holotype of Gnatholepis scapulostigma, 1(36.5), Engebi Island, Enewatak, A.D. Welander, 11 August 1949. AMS
I.37692-002, 1(36.5), Eniwetok Atoll, R. Nolan, 28 May 1972. HAWAII: BPBM 37847, holotype of Gnatholepis
cauerensis hawaiiensis, 1(35), N shore off Pupukea, Oahu, R. Holcom, August 1997. BPBM 37861, 3(14–38.5),
Kona coast off airport, Hawaii, J. Randall, 22 September 1997; BPBM 15131, 13(24–42), Baldwin Packer’s
property, Maui, W. Gosline and E. Hunter, 5 August 1955. SOCIETY ISLANDS: CAS 51602, 5(26–36), W side
Papetoai Bay, Moorea, 18 September 1956; CAS 51528, 1(24.5), inside Teputo Pass, Papeari District, Tahiti, 4 July
1957; BPBM 8108, 2(24.5–35), Popote Bay, Papara, Tahiti, 13 March 1969. MNHN 1980-94, 1(32.5), Gambier
Islands, P. Fourmanoir; AMS I.22176-003, 4(33–37), Ahahu Bay, Nuku Hiva, 3 September 1976; BPBM 11869,
8(21–53), N end Hanau Bay, Fatu Hiva, Marquesas, J. Randall, D. Cannoy, R. McNair, 21 April 1971; BPBM
11003, 2(29.5–42), W side Ua Huka, N of Mt Takatai, Marquesas, J. Randall, J. Haywood, R. McNair, 7 May 1971;
BPBM 12621, 3(28–39), S end Akuoa Bay, Nuku Hiva, Marquesas, J. Randall, D. Bryant, R. McNair, 12 May
1971; BPBM 10398, 1(330, Hevane Bay, Hana, Tahuata, Marquesas, 16 October 1961; BPBM 12764, Nuku Hiva,
Marquesas, 16 May 1971; BPBM 17276, holotype of Gnatholepis cauerensis australis, 41 mm SL, entrance to
Haurei Bay, Rapa, Austral Islands, J. Randall and D. Cannoy, 10 February 1971; BBPBM 38376, 1(35.5), entrance
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to Haurei Bay, Rapa, Austral Islands, J. Randall and D. Cannoy, 10 February 1971; BBPBM 17319, 1(35.5),
entrance to Haurei Bay, Rapa, Austral Islands, J. Randall and D. Cannoy, 14 February 1971; BPBM 17267, 1(38), E
side Akatamiro Bay, Rapa, Austral Islands, J. Randall and D. Cannoy, 8 February 1971; USNM 379839,
2(38.5–40), Anarua Bay, Rapa, Austral Islands, J.T. Williams and party, 7 November 2002; USNM 379757,
4(19.5–33.5), NW of Isle Tautoroa, Baie Anatkurinako, Rapa, Austral Islands, J.T. Williams and party, 13
November 2002; USNM 379741, 5(20.5–37.5), N coast of Point Kauira, Rapa, Austral Islands, J.T. Williams, 15
November 2002; USNM 379787, 2(23–31), S of Isle Tauna at mouth of Haurei Bay, Rapa, Austral Islands, J.T.
Williams and S. Planes, 15 November 2002; USNM 379718, 3(35–39), N side channel, Haurei Bay, Rapa, Austral
Islands, J.T. Williams and party, 19 November 2002; USNM 379630, 3(35–39), N end of Point Tematapea, Rapa,
Austral Islands, J.T. Williams, 19 November 2002. PITCAIRN ISLANDS: BPBM 16846, 2(38.5–41.5), off Bounty
Bay, Pitcairn, J. Randall and party, 26 December 1970.
FIGURE 14. Gnatholepis cauerensis, Flores, Indonesia. Photo by Rudie Kuiter.
Other material; no data taken. YEMEN: SAIAB 69164, 1(36), 12° 44’ N 14° 42’ E, Ras Imram, Gulf of
Aden, south Yemen, P. and U. Zajonz, 24 March 1998. SOUTH AFRICA: SAIAB 53491, 1, Tshani, Transkei;
SAIAB 74853, 1, Eelskin, Kwa-Zulu Natal; SAIAB 46430, 3, Aliwal Shoal, Kwa-Zulu Natal; SAIAB 69382, 3,
Aliwal Shoal, Kwa-Zulu Natal. MOZAMBIQUE: SAIAB 74506, 2, Inhambane. ALDABRA: USNM 32547, 2,
Aldabra Atoll. TANZANIA: BPBM 16392, 1, Chole Bay, Chole Island, Mafia Island. SEYCHELLES: USNM
327484, 29, St Joseph Atoll, Amirantes Islands. MAURITIUS: SAIAB 57882, 9, Flic en Flac; SAIAB 52054, 3,
Albion Fisheries Research Centre; USNM 308208, 21, Agalega Islands. MALDIVES: BPBM 34390, 4, Furana
Island, North Malé Island; BPBM 34574, 2, South Malé Island. PHILIPPINES: AMS I.21922-009, 4, Caban
Island, Batangas Province; AMS I.21908-002, 8, Sombrero Island, Batangas Province; USNM 261671, 8, Negros
Oriental. INDONESIA: BPBM 32249, 1, Tulamben, Bali; AMS I.34501-018, 1, Waliti, Sao Wisata Resort, Flores;
NSMT-P.71338, 4, Terawangan Island, Lombok; NSMT-P.71354, 18, Ambon Bay, Ambon; NSMT-P.64522, 6,
Kotania Bay, Seram; BPBM 37362, 1, Kebola Bay, Alor; BPBM 36644, 1, Nil Desperandum Reef, Banda Sea;
BPBM 37641, 1, Sarabua Bay, Siberut Island, Mentawi Islands. TAIWAN: SAIAB 31091, 4, Kenting National
Park. JAPAN: AMS I.27366-002, 1, Amitori Bay, Iriomote-jima. JAPAN: BPBM 15021, 2, Ishigaki-jima, Ryukyu
Islands. MICRONESIA: USNM 224765, 12, Ponape, Caroline Islands. PAPUA NEW GUINEA: USNM 327485,
5, Madang Harbour. AUSTRALIA: AMS I.28992-009, 1, Horsburgh Island, Cocos-Keeling Islands. NTM
S.12883-015, 15, Cartier Reef, Timor Sea, Western Australia; NTM S.11061-007, 3, One Tree Island. NEW
CALEDONIA: MNHN 1980-158, 3; MNHN 1980-124, 1, Ouvea, Loyalty Islands. VANUATU: AMS I.39010-
100, 1, Santa Cruz; USNM 380364, 2. FIJI: USNM 243207, 9, Viwa Island. MARSHALL ISLANDS: AMS
I.37709-003, 1, Eniwetok; USNM 368817, 16, Bikini Atoll. TONGA: USNM 340172, 16; USNM 340191, 6;
USNM 340185, 3. SOCIETY ISLANDS: ex-MNHN 1984-168, 3, Tiahura, Moorea.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
FIGURE 15. Gnatholepis cauerensis over dark sand, Tulamben, Bali, Indonesia. Photo by Gerry Allen.
Description. Based on 117 specimens, 24.5–56 mm SL. An asterisk indicates the counts of the holotype of
Gobius cauerensis.
First dorsal VI*; second dorsal I,10–I,11* (mean I,11.1); anal I,10–12 (mean I,11.2, 11 in holotype), pectoral
rays 12–18 (mean 16.3; 16 on right side, 17 on left in holotype), segmented caudal rays nearly always 17*; caudal
ray pattern nearly always 9/8*; branched caudal rays 6/5 to 8/7 (usually 7/6, 7/7 in holotype); lateral scale count
24–30 (mean 26.7, 25 in holotype); TRB 8½–12 (mean 10.0, 9 in holotype); predorsal scales 7–12 (mean 8.8, 10 in
holotype); circumpeduncular scales 11–13 (mean 12.2). Gill rakers on outer face of first arch 1–2 + 3–4 (in 25,
usually 1+4).
Body compressed, width at anus 10.4–15.5% (mean 13.1%) of SL. Body rather slender in appearance, body
depth at anus 13.7–24.9% (mean 21.6%) of SL, body depth at first dorsal fin origin 18.3–24.7% (mean 22.2%) of
SL. Head compressed, slightly broader ventrally, slightly deeper than wide, HL 24.1–30.7% (mean 27.4%) of SL;
head depth at posterior preopercular margin 62.0–81.0% (mean 68.6%) of HL; head width at posterior preopercular
margin 57.8–78.5% (mean 67.5%) of HL; head profile bluntly pointed to rounded; nape profile low, slightly
convex. Mouth subterminal to nearly terminal, slightly oblique; jaws generally reaching to below anterior half of
eye to just beyond front of eye; upper jaw length 27.1–40.5% (mean 34.6%) of HL. Upper lip smooth, narrower
than lower, lower lip with papillose ridge close to teeth, with twist or fold posteriorly, forming triangular flap, lip
narrowly interrupted at chin. Eye moderate to relatively small, dorsolateral, 20.1–31.3% (mean 26.8%) of HL;
preorbital width 16.1–31.6% (mean 20.1%) of HL. Snout bluntly pointed to rounded, 21.1–39.0% (mean 32.2%) of
HL; posterior naris round to almost triangular, close to anterior margin at middle of eye; anterior naris in short tube,
higher on posterior margin of eye, about level with middle of eye or somewhat ventral to it. Interorbital narrow,
5.3–14.3% (mean 7.6%) of HL. Caudal peduncle compressed, length 11.4–23.4% (mean 16.2%) of SL; caudal
peduncle depth 9.1–17.5% (mean 11.7%) of SL.
First dorsal fin rounded to roughly triangular, with no spines elongate; third to fourth dorsal spine longest;
when adpressed, spine tips reaching to first to third element of second dorsal fin. Second dorsal spine length
15.1–21.7% (mean 17.3%) of SL; third dorsal spine length 13.1–22.2% (mean 17.6%) of SL. Second dorsal fin as
tall as first dorsal fin, posteriormost rays usually longer than anterior, fin pointed to slightly rounded posteriorly.
Anal fin lower than second dorsal fin, anteriormost rays shorter than posterior few rays; fin pointed to slightly
rounded posteriorly. Second dorsal and anal fin rays, when adpressed, just reaching caudal fin rays. Pectoral fin
oval to somewhat pointed, central rays longest, 18.6–28.6% (mean 25.7%) of SL; fin reaching back to above first
few anal fin elements. Pelvic fins fused, frenum with distinctive finely fimbriate margin, fins round to somewhat
oval, reaching to first anal fin elements, 22.9–31.5% (mean 26.9%) in SL. Caudal fin moderate, oval to rounded,
23.7–34.3% (mean 30.5%) of SL.
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Gill opening restricted, extending anteriorly to lower edge of pectoral base or to just under opercle. Up to three
slender gill rakers on outer face of first arch, close to angle of arch, remaining few rakers very short, pointed;
pointed papillae present along inner face of first arch; outer rakers on second gill arch consisting of two series of
papillae, one low and one (mostly anterior) of slightly pointed papillae; outer rakers on remaining arches low,
stubby, broad-based. Inner face of upper limb of first gill arch, and to lesser extent, upper limbs of other arches,
covered with low dense fleshy papillae, usually forming clumps or groups; dorsal portion of arch may have short
fleshy protuberances ending in one or several papillae; inner face of second arch smooth, without papillae but with
series of short fleshy rakers with rounded, finely papillose tips. About one quarter to one-third of first gill arch
bound by membrane to inner face of opercle. Tongue short, tip bilobed to concave.
Teeth in upper jaw in two to three rows across front and one row at side of jaw, outermost row teeth largest,
curved and pointed, largest teeth at front of jaw on either side of symphysis (smaller in females); innermost row
teeth very small, sharp and evenly sized. Teeth in lower jaw in two to three rows, arranged similarly to upper jaw
but outer row teeth may be smaller (in females); posteriormost one to three outer row teeth enlarged to somewhat
recurved in males.
Predorsal scales cycloid, ctenoid body scales usually reaching forward to rear corner of opercle to just over
opercle, occasionally ctenoid scales reaching forward to above rear preopercular margin; nape midline scales
always cycloid (Fig. 4C–D). Opercle covered with cycloid scales, a few ctenoid scales may be present.
Preopercular scales cycloid, occasionally extending anterior to vertical dark cheek-bar below eye. Breast with
cycloid scales reaching up to below mid-opercle or to below rear half of opercle. Pectoral fin base covered with
cycloid scales; occasionally with some ctenoid scales. Belly scales along midline usually cycloid; ctenoid scales
may be present posteriorly.
Head pores as in G. a n j e re n s i s (Fig. 1), as are the sensory papillae.
FIGURE 16. Gnatholepis cauerensis, from Sodwana Bay, Kwa-Zulu Natal, South Africa, showing great similarity to G.
thompsoni. Photo by Phil Heemstra.
Coloration of fresh material. Good illustrations of living fish are shown in Senou et al. (2004: 242), Hayashi
and Shiratori (2003: 97, as G. s c a p u l o s t i g m a ), Kuiter and Tonozuka (2001: 673, as G. c a u e re n s i s and G.
scapulostigma), Myers (1991: pl. 124B, C; 1999: pl. 161E), while fresh-dead specimens are shown in Randall and
Greenfield (2001: pl. I, G, H; pl.II, A–D). Anderson (2005: 97) shows the western Indian Ocean spotted form from
the Maldives.
The range of colouring shown in these publications, and others, shows that there is considerable variation in
live colouring in G. c a u e r e n s i s . Live fishes photographed on dark sand are the most intensely coloured (e.g.
Randall & Greenfield 2007: 306, fig. 2). In our experience, determining what is “typical” for the species is difficult
but the upper fish illustrated in Senou et al. (2004: 242) comes close (and see Fig. 13).
Gnatholepis cauerensis usually has the body translucent white with five to seven rows of small variably
coloured (dull orange, brown, red-brown to blackish) spots and/or thin lines and about six indistinct diffuse orange-
brown to purple-brown blotches along mid-side and the last much smaller, similarly coloured mark may be present
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
on base of caudal fin; narrow vertical black line on top of each eye, lines may join each other across interorbital
space, and narrow black to reddish brown vertical bar crossing cheek below eye; small dark mark of variable shape
(often broad U- or W-shaped) above pectoral fin base, with small yellow spot in centre; side of head with small
purple-brown to reddish marks and blotches that often form a cluster on either side of the dark vertical bar crossing
the cheek from below the eye; dorsal and anal fins transparent with rows of indistinct dusky fine streaks or spots;
anal fin translucent pearly whitish; pectoral fin transparent with white speckling on membranes basally; pelvic fin
translucent pale white; caudal fin transparent with translucent white rays with rows of thin streaks, lines or fine
spots along membranes or edges of rays (Figs 13–16).
FIGURE 17. Gnatholepis cauerensis over dark sand, Normanby Island, Papua New Guinea. Photo by Gerry Allen.
Coloration of preserved material. Based on typically coloured specimens from Ceram (variations discussed
below). Head and body yellowish white to whitish with up to seven variably developed thin brownish lines running
along length of body; usually two most conspicuous lines (lines number 5 to 6 if counting down from dorsum)
running from behind pectoral fin base, this pair of lines overlying six square to rectangular brownish blotches
(these blotches may be underlain by five to six vertically oval, faintly brownish bars similar to those in G.
anjerensis); next most conspicuous pair of lines (numbers 2 to 3) running from just above pectoral fin base and
becoming indistinct on upper part of caudal peduncle, this pair of lines enclosing a series of up to 10 diffuse
brownish square blotches, usually more conspicuous on anterior part of body. Dorsum crossed by about nine faint
dusky square saddles (first saddle on nape before first dorsal fin), which may coalesce with the square blotches
along upper part of body.
Predorsal, snout and side of head faintly blotched with pale brown; blotches along predorsal often paired.
Opercle with (often indistinct) pale brown line from centre of preopercular margin obliquely back to upper half of
opercle rear margin; line may curve up to rear corner of opercle. Narrow blackish, light brown to faint grey cheek-
bar from ventral edge of eye running vertically to slightly obliquely down to end on lower preopercular edge, but
not extending onto branchiostegal membranes. Dense blackish to dark brown line from middle to rear half of iris
running over top of eye (line may be slightly oblique), may meet its counterpart in interorbital space. Upper lip
whitish, crossed by short brownish bar (starting from below anterior naris) as in G. a n j e re n s i s ; similar brownish bar
or line at middle of lip. Cheek with diffuse brownish horizontal line crossing vertical cheek-bar; brownish line
variable in shape and intensity, its pigment often surrounding one or two round pale spots. Above pectoral fin base,
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brownish to dark brown, roughly U-shaped flat-based blotch surrounding small very pale brown round spot; brown
blotch often partly connected to brown line (may be partly broken-up or indistinct) running from behind eye along
top of preopercle; blotch usually paler in small specimens. Pectoral fin base with horizontal light brown line
crossing just above middle of fin base, line usually extending onto lower part of fin.
First dorsal fin transparent, with about three thin wavy light brown horizontal lines beginning at first spine;
lowermost one or two lines darkest, especially anteriorly; distal part of fin may be plain pale brownish. Second
dorsal fin with similar few thin light brown horizontal lines which, after the first few fin rays, transform into plain
brown pigment or series of vertical dark streaks on membrane between fin rays; in some specimens fin almost
unpigmented. Anal fin plain dusky to brownish, may be darker distally. Caudal fin transparent to translucent
brownish, sometimes with pale brown streaks or rows of fine brown spots along fin membranes, especially near fin
base. Pectoral fins transparent to translucent dusky. Pelvic fins and frenum translucent plain pale brownish, may be
darker between fourth and fifth rays.
Var ia tio n . A number of specimens from a range of localities (Kwa-Zulu Natal to the Philippines to the
Marquesas) are more heavily marked (though often from white-sand habitats), with all markings darker (Fig. 15).
The seven lines along the body are intensified into a series of dark brown partly connected spots, the six
rectangular blotches along the mid-side of the body are more prominent purplish brown, the thin lines along both
dorsal fins are dark brown to blackish anteriorly and are broken into series of spots; those along the second dorsal
fin tending to form short longitudinal streaks with none of the vertical dark lines between rays. The caudal fin is
crossed by many rows of small dark spots, which are slightly larger and darker toward the fin base.
In particular, photographs of specimens from western Indian Ocean locations such as Kwa-Zulu Natal coast,
South Africa, Mauritius and Maldives all show a similar spotted pattern (Fig. 16). These fish are almost identical in
colour pattern to G. t h o m p s o n i .
Specimens from Rapa (which show lines, not rows of spots, along the body) show a similar pattern on the
caudal fin: thin light brown lines following the fin rays, darker toward the centre and base of the fin. This pattern is
also found in specimens from Hawaii (which Greenfield & Randall (2001) placed in subspecies G. c . h a w a i i e n s i s ).
Among the largest specimens examined, from Taiwan and Middleton Reef, specimens have the six or seven
horizontal lines along the body considerably darker, with the two to three lines closest to the mid-side of the body
partly extending on to the caudal fin. One Middleton Reef specimen is heavily marked with dark lines along the
body. Many Hawaiian specimens are similarly strongly marked (some labelled as being from black sand habitats),
having six or seven dark brown lines along the body. This pattern is also seen in photographs of living fish from
localities such as Izu Peninsula (Japan) and Normanby Island (New Guinea) (Fig. 17).
FIGURE 18. Gnatholepis cauerensis, smaller specimen is holotype of Gobius cauerensis, RMNH 4523. Photograph by Doug Hoese.
Distribution. A wide-spread Indo-Pacific species, known from South Africa to Pitcairn Island, in the Pacific
north to Miyake-jima, Japan and south to Sydney Harbour, Australia, and in the Indian Ocean north to Djibouti
(Gulf of Tadjourah) and south to Aliwal Shoal, Kwa-Zulu Natal.
Ecology. Recorded at depths of 1–33 m, over sand near live or dead corals, to sand flats near rocky reef.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Comparisons. This species has frequently been confused with G. a n j e r e n s i s but can be distinguished from it
by having cycloid predorsal scales (versus ctenoid predorsal scales in G. a n j e r e n s i s ); its smaller size and generally
slender appearance (up to 56 mm SL versus 84 mm SL and generally stocky appearance of G. a n j e re n s i s ); having a
generally rounded to roughly triangular first dorsal fin with third or fourth dorsal spine longest (versus roughly
square in shape, with fifth dorsal spine nearly as long as third to fourth spines in G. a n j e re n s i s ); body pale with five
to seven rows of small orange, brown, red-brown to blackish spots and/or thin lines and about six indistinct diffuse
orange-brown to purple-brown blotches along mid-side (versus pale body covered with irregular rows of small
blackish to reddish spots and five to six diffuse dusky blotches along the lower mid-side of the body, and blue spots
on side of head and anterior part of body in G. a n j e r e n s i s ); and small dark mark, often W-shaped, above pectoral fin
base, with small yellow spot in centre (versus small round orange to gold spot, often with short dark line
underneath, in G. a n j e r e n s i s ).
Remarks. The holotype of Gobius cauerensis Bleeker, 1853, is one of two specimens tied together in RMNH
4523 (31.2–32.6 mm SL; 38.9–39.9 mm TL), with its size given in the original description as 39 mm TL; the
second smaller specimen is thus the holotype (Fig. 18). Randall and Greenfield (2001) give the holotype number as
RMNH 6523.
Gnatholepis scapulostigma Herre, 1953, was described from the type (and only specimen), a male 36.5 mm
long, but Herre did not state where the type was deposited. Randall and Greenfield (2001) found it in the USNM
main collection (USNM 344429), not labelled as a type but agreeing otherwise with data in Herre’s description.
They state the fish has 10 anal fin rays and 17 pectoral fin rays; we obtained counts of 11 anal rays and 16 pectoral.
Gnatholepis inconsequens Whitley, 1958, was described from Heron Island on the southern Great Barrier
Reef. Randall and Greenfield (2001) placed this as a synonym of G. c a u e r e n s i s based on pectoral fin ray count.
Other specimens from the area, including AMS material collected at the same time as the type, are G. a n j e re n s i s .
Although the specimen is stained green (from “copper-stained formalin” (Whitley 1958)), the typical G. c a u e re n s i s
dark shoulder blotch can be discerned and the pectoral fin lacks the speckling of melanophores as found in G.
anjerensis (all predorsal scales are missing).
Wass (1984) briefly described a pale Gnatholepis from Samoa (AMS I.22003-001); his description agrees with
G. c a u e re n s i s . Many records of G. a n j e re n s i s in the literature before the 1990s refer to G. c a u e re n s i s and vice versa,
due to confusion over the use of these two names (see Myers 1991, 1999).
Gnatholepis cauerensis australis was described as a subspecies by Randall and Greenfield (2001) from
Haurei Bay, Rapa, French Polynesia (holotype BPBM). This material was first tentatively identified as G.
anjerensis by Randall et al. (1990b). Randall (2009: 180–181) brought this to full species status, based on its
pectoral fin ray counts of 17–19 (modally 19), compared to G. c a u er en si s’ pectoral ray counts of 16–19 (modally
17), based on 10 BPBM specimens (Randall included counts from both sides in his Table 1 of 2009). We found
pectoral ray counts of 14–19 (modally 18), based on 19 Rapa specimens from BPBM and USNM. A difference
in adult size between G. a u s t ra l i s (sensu Randall 2009) and G. c a ue re n s i s is remarked upon by Randall (2009),
who cites 40 mm SL as the largest G. c a ue re n s is out of 80 specimens that he had examined, with only two being
over 38 mm SL, while the 31 specimens of G. a u st r a li s he looked at included eight over 38 mm SL, with the
largest being 49 mm SL. Of the 117 G. c a u ere ns i s we examined, the largest specimen was 56 mm SL (from
Marquesas), with other specimens over 50 mm SL from Taiwan, Middleton Reef and Elizabeth Reef. The modal
SL for all G. ca u ere n si s was 39 mm, while mean SL was 37.8 mm. Of our 19 specimens from Rapa and Pitcairn,
the maximum size examined was 41.5 mm SL, a mode of 35.5 mm and a mean of 35.7 mm SL. We leave G.
australis as a synonym of G. c a u e re n s is .
Randall and Greenfield (2001) described the subspecies Gnatholepis cauerensis hawaiiensis, from off
Pupukea, Oahu, Hawaiian Islands (holotype BPBM 37847). They separated this subspecies from G. c a u e re n s i s
cauerensis by its having a shorter caudal peduncle (in head length), in the caudal fin having a dark line along the
membrane between rays instead of dark spots and in having small blue spots on the lower half of the body. Later
Randall (2009) elevated the Hawaiian form to full species, G. h a w a i i e n s i s , on the basis of caudal peduncle length:
1.7–1.9 (in head length) in G. c a u e r e n s i s and 1.5–1.7 in G. h a w a i i e n s i s . We were unable to determine a consistent
difference in caudal peduncle length between any Gnatholepis species (mean length of 15.7–17.0% in SL). The
lines, streaks and rows of dark spots on the caudal fin in G. c a u e re n s i s vary greatly, with specimens from localities
such as Ogasawara Islands (Japan), Cartier Reef and Shark Bay (Western Australia), One Tree Island (Queensland)
and Fiji showing these lines as described for G. c a u e re n s i s h a w a i i e n s i s . The small pale blue spots on the lower part
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of the head and body vary also (more heavily pigmented specimens more likely to show these pale spots)—and are
usually only discernible in photographs of living or recently dead fish. Therefore we do not agree with Randall
(2009: 181) that the Hawaiian form is a separate species.
TABLE 11. Measurements of specimens of Gnatholepis cauerensis, expressed as percentage of standard length (SL) or
head length (HL); n = 112.
One specimen from Ras Imran in the Gulf of Aden, Yemen (SAIAB 69164) resembles G. c a u d i m a c u l a t a but
lacks the vertical dark blotch on the caudal base (has the faint short horizontal mark typical of G. c a u e r e n s i s ) and
has a straight (not curved) black line atop the eyeball. This specimen is the “farthest north” that we have examined
for G. c a u e r e n s i s .
Randall (in litt. to HKL) considered that G. c a u e re n s i s specimens from Marquesas constituted an undescribed
species, due to differences in colour and pectoral fin ray count. In Randall (1985), Marquesas specimens are
identified as G. a n j e re n s i s ; in Randall and Earle (2000) they are identified as G. c a u e re n s i s . One of the fish (one of
three in BPBM 12621) is shown in Plate IIA of Randall and Greenfield (2001). Five Marquesas specimens (in
BPBM 12621, 11869, 11003) had counts of 17 pectoral fin rays and five had 18. A SAIAB specimen photographed
from Mauritius (not examined) and a wild fish from Bali, Indonesia, photographed by Rudie Kuiter (not collected)
have similar heavy spotting to the Marquesas fish, with the adjoining lines hardly visible and only five lateral
blotches present, not six. The variation in these fishes gives one much to think about.
It is of concern that some specimens identified by Randall and Greenfield in 2001 as G. a n j e r e n s i s , were
subsequently identified by us as G. c a u e re n s i s . These include fish from Heron Island and Middleton Reef,
Queensland, and may be some of the “exceptional” specimens from the southern Great Barrier Reef that they
mention as having distinct longitudinal lines on the body (Randall & Greenfield 2001: 5). Many specimens and
photographs from Japanese waters show fish with distinct longitudinal lines along the body (e.g. Hayashi &
Shiratori 2003; Senou et al. 2004).
Holotype
(male)
Means
(males)
Maximum
(males)
Minimum
(males)
Means
(females)
Maximum
(females)
Minimum
(females)
Standard length 31.7 37.2 56.0 23.0 36.3 51.0 27.0
Head length in SL 26.5 27.2 30.7 24.1 27.9 29.8 24.9
Head depth in HL 81.0 69.4 81.0 62.9 67.2 77.6 62.0
Head width in HL 67.9 69.0 78.5 58.5 66.8 76.6 57.8
Body depth at anus in SL 24.9 21.7 24.9 18.4 21.8 24.7 13.7
Body depth at D1 origin in SL – 22.0 23.8 18.6 22.7 24.7 18.3
Caud. ped. length in SL 18.3 16.5 23.4 12.5 16.1 20.9 11.4
Caud. ped. depth in SL 12.3 11.8 12.9 9.1 11.8 17.5 10.8
Snout length in HL 31.0 33.2 39.0 21.1 31.4 38.1 25.9
Eye width in HL 29.8 26.6 29.9 20.1 26.8 31.3 23.7
Jaw length in HL 40.5 35.0 40.5 27.8 33.8 40.2 27.1
Interorbital in HL 14.3 7.8 14.3 5.3 7.6 9.9 6.0
Preorbital width in HL – 20.2 23.7 16.1 20.2 31.6 16.3
Pectoral fin in SL 18.6 25.5 27.4 18.6 26.0 28.6 22.5
Pelvic fin in SL 28.4 27.3 31.5 23.9 26.5 30.6 22.9
Caudal fin in SL 23.7 31.2 34.3 23.7 29.2 32.3 24.0
Adpressed D1 in SL – 24.4 31.8 15.5 23.0 26.3 16.7
3
rd
D1 fin spine in SL – 17.7 22.2 13.1 17.4 19.7 14.9
4
th
D1 fin spine in SL – 17.9 20.5 13.1 16.9 19.5 14.2
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
In Thacker (2004a: 576), Figure 2 shows a strict consensus tree with three unlabelled outliers grouped together
at the base of her G. s c a p u l o s t i g m a (= G. c a u e r e ns i s ) clade. These were two specimens from Moorea and one from
Aitutaki, Cook Islands, and there was no morphological information to indicate that they were different (Thacker,
in litt. 27/1/06). Other specimens from the same two localities grouped with her G. k n i g h t i clade (but separately
from material from Hawaii).
Gnatholepis gymnocara Randall and Greenfield, 2001
(Table 5)
?Gobius pauper de Vis, 1884: 687 (Moreton Bay).
Gnatholepis gymnocara Randall and Greenfield, 2001: 14 (Supply Bay, 22°12.24'S, 150°28.53"E, Townshend I., Queensland).
Diagnosis. (modified from Randall & Greenfield 2001) A small Gnatholepis (up to 35.4 mm SL) lacking scales on
side of cheek, opercle and predorsal midline, few cycloid scales present on pectoral fin base and on breast; no
distinct flap on end of lower lip (low fold may be present); no distinct canine or enlarged teeth; second dorsal fin
rays I,11; anal fin rays I,11–12 (usually I,12); pectoral rays 15–19 (usually 17); lateral scales 26–28, ctenoid;
predorsal scales absent from nape midline, scales on side of head reaching to nearly behind eyes; body pale with
dusky speckling and mid-lateral row of six dusky blotches and horizontal row of darker spots and short streaks, an
irregular cross-shaped dark mark on cheek extending down from eye, pupil-sized brown spot just above mid-point
of caudal fin base, males with small black spot on rear part of first dorsal fin.
Material Examined. AMS I.34318-032, paratypes of Gnatholepis gymnocara, 32(17–33), Supply Bay,
Townshend Island, Queensland, S. Reader and party, 16 September 1998.
Remarks. In their description of G. g y m n o c a r a , Randall and Greenfield included two paratype specimens from
the Northern Territory (AMS I.23930-011). As no other specimens of G. g y m n o c a r a have so far been found in the
Territory, these paratypes were examined and were found to be female G. a r gu s (the larger specimen had a second
dorsal fin count of I,11, but anal ray count of I,11, while the smaller specimen had I,10 second dorsal rays and I,11
anal rays; both have scaleless pectoral fin bases and typical female G. a rg u s colouring).
The species described as Gobius pauper De Vis, 1884, from Moreton Bay, Queensland, has no known type
specimen. It is possible that it may have been deposited in the Queensland Museum, but it is not there now and its
whereabouts are unknown. From De Vis’ original description, it appears to be a species of Gnatholepis, as noted by
Hoese and Larson (2006). De Vis described his species as being without canines, which implies G. g y m n o c a r a (and
not G. c a u e r e n s i s ), but we are reluctant to synonymise G. p a u p e r with G. g y m n o c a r a or any other species without
more certainty as to the identity of the type specimen(s).
Gnatholepis knighti Jordan and Evermann, 1903
(Figs 12D, 19–20, Tables 5–8, 12)
Gnatholepis knighti Jordan and Evermann, 1903: 204 (Hilo, Hawaii).—Jordan and Evermann 1905: 11 (coral reefs at Oahu);
Jordan and Evermann 1926: 12 (Hawaii); Koumans 1940: 133, 155; Böhlke 1953: 112; Böhlke 1984: 107; Ibarra and
Stewart 1987: 40; (in part); Thacker 2004b: 578–579.
Gnatholepis anjerensis—Randall et al. 1985: 70 (Johnston Island); Randall and Greenfield 2001: 3, 5–6 (in part); Greenfield
and Randall 2004: 524–525 (Kane’ohe Bay, O’ahu).
Diagnosis. A large stocky Gnatholepis (up to 64 mm SL) with cycloid scales on predorsal region, cheek and
pectoral fin base, distinct flap present at end of lower lip; at least three dark spots along first spine of first and
second dorsal fins, followed by at least three rows of dark spots or short streaks, pectoral fin plain translucent to
pale brownish; transverse black line crossing upper part of eye usually split in two; anal fin plain dusky, often with
one to several blackish rounded to oval spots posteriorly, or with dark streaks following membrane; second dorsal
and anal fin rays nearly always I,11; pectoral rays 15–17, usually 16; lateral scales 25–29, usually 27, 7–10
predorsal scales 7–10, usually 8–9.
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FIGURE 19. Gnatholepis knighti, 2 m depth, Kaneohe Bay, Hawaii. Photo by Rob Myers.
Material Examined. NORTH-WESTERN HAWAIIAN ISLANDS: BPBM 34873, 5(29–36), Midway Atoll,
wreckage at shore between cargo pier and launching ramp, J.E. Randall and J.L. Earle, 18 September 1991.
HAWAII: Paratypes of Gnatholepis knighti, CAS-SU 7468, 7(32.5–43), Hilo, D.S. Jordan and B.W. Evermann,
1901–1902. Paratypes of Gnatholepis knighti, USNM 144094, 5(30.5–43), Hilo, D.S. Jordan and B.W. Evermann,
1901. Paratype of Gnatholepis knighti, BPBM 1699, 1(34), Hilo, U.S. Fish Commission, 1901. USNM 78072,
23(24.5–44), Hawaii, U.S. Fish Commission, 1901; BPBM 19665, 10(58–65), SW side Oahu, Campbell Industrial
Park, B. Carlson, J. McKinney and A. Suzumoto, 11 September 1975; BPBM 28720, 1(42.5), South Kahala,
Waiulua Bay, J.E. and H.A. Randall, 11 August 1982. LINE ISLANDS: BPBM 34045, 3(27–45), Johnston Atoll,
R.K. Kosaki and party, 25 July 1989.
Description. Based on 28 specimens, 32.5–64.0 mm SL.
First dorsal VI–VII (usually VI); second dorsal I,10–I,12 (mean I,10.9); anal I,10–11 (mean I,10.9), pectoral
rays 15–17 (mean 16.1), segmented caudal rays nearly always 17; caudal ray pattern usually 9/8; branched caudal
rays 6/7 to 8/7 (usually 7/6); lateral scale count 25–29 (mean 27.1); TRB 8–11½ (mean 10.3); predorsal scales 7–10
(mean 8.7); circumpeduncular scales always 12. Gill rakers on outer face of first arch 1+4 (in 2).
Body compressed, width at anus 11.4–15.7% (mean 13.2%) of SL. Body rather stocky in appearance, body
depth at anus 20.6–26.0% (mean 23.2%) of SL, body depth at first dorsal fin origin 21.0–26.1% (mean 23.8%) of
SL. Head compressed, broader ventrally, slightly deeper than wide, HL 25.0–29.9% (mean 27.3%) of SL; head
depth at posterior preopercular margin 66.3–85.6% (mean 74.5%) of HL; head width at posterior preopercular
margin 61.2–80.0% (mean 70.9%) of HL; head profile bluntly pointed; nape profile low, relatively straight. Mouth
subterminal to nearly terminal, slightly oblique; jaws generally reaching to below anterior margin of eye; upper jaw
length 29.4–40.6% (mean 35.3%) of HL. Upper lip smooth, narrower than lower, lower lip papillose close to teeth,
with twist or fold posteriorly, forming triangular flap, lip interrupted at chin. Eye moderate to relatively small in
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
large specimens, dorsolateral, 24.8–32.0% (mean 27.6%) of HL; preorbital width 19.1–27.9% (mean 22.5%) of
HL. Snout bluntly pointed, 28.6–44.0% (mean 36.4%) of HL; posterior naris round to almost triangular, close to
anterior margin of eye; anterior naris in short tube, higher on posterior margin of eye, about level with middle of
eye or somewhat ventral to it. Interorbital narrow, 6.1–10.4% (mean 8.6%) of HL. Caudal peduncle compressed,
length 15.2–19.0% (mean 17.0%) of SL; caudal peduncle depth 10.2–13.5% (mean 11.9%) of SL.
First dorsal fin rounded to almost square, with no spines greatly elongate; first, second or third spine longest;
when adpressed, spine tips reaching to first or second element of second dorsal fin. First spine length 14.3–25.6%
(mean 19.3%) of SL; second dorsal spine length 14.7–20.2% (mean 16.1%) of SL; third dorsal spine length
14.5–18.4% (mean 16.3%) of SL. Second dorsal fin as tall as first dorsal fin, rays longer anteriorly than posteriorly,
fin pointed to slightly rounded posteriorly. Anal fin lower than second dorsal fin, anteriormost rays shorter than
posterior few rays; fin pointed to slightly rounded posteriorly. Second dorsal and anal fin rays, when adpressed,
usually reaching caudal fin. Pectoral fin rounded to slightly pointed, central rays longest, 23.1–28.5% (mean
25.8%) of SL; fin reaching back to above anal fin spine or first few anal fin elements. Pelvic fins fused, frenum
with distinctive finely fimbriate margin, fins round to somewhat oval, reaching to first few anal fin rays,
22.6–29.6% (mean 25.9%) in SL. Caudal fin moderate, oval to rounded, 26.1–35.6% (mean 29.7%) of SL.
Gill opening restricted, extending anteriorly to lower edge of pectoral fin base. Gill raker form generally as in
G. a n j e re n s i s ; about a quarter of first gill arch bound by membrane to inner face of opercle. Tongue bilobed.
Teeth as in G. a n j e re n s i s .
Predorsal scales cycloid, ctenoid scales on body reaching over pectoral fin base, sometimes reaching onto side
of nape over opercle, nape midline always with cycloid scales. Opercle with cycloid scales, upper or central part
occasionally with ctenoid scales. Preopercular scales cycloid, which may extend anterior to vertical dark cheek bar
below eye. Breast with cycloid scales, extending forward nearly to rear edge of preopercle. Pectoral fin base scales
cycloid. Belly scales usually cycloid; ctenoid scales present on posterior half of belly midline in some specimens.
FIGURE 20. Gnatholepis knighti, 2 m depth, Kaneohe Bay, Hawaii. Photo by Rob Myers.
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Head pores and sensory papillae as in G. a n j e re n s i s (Fig. 1).
Coloration of fresh material. This species has been described and illustrated in Greenfield and Randall
(2004: 524–526, fig. 64); the dark spots on the white anal fin are visible in this figure. A photograph by Jack
Randall of a large dead specimen from Oahu resembles G. a n j e r e n s i s but the pectoral fin is translucent with fine
whitish speckles on the fin rays; the anal fin is relatively plain dusky white, and the dorsal part of the body has
scattered small dense black spots (and see Figs 19–20 from Kaneohe Bay).
TABLE 12. Measurements of specimens of Gnatholepis knighti, expressed as percentage of standard length (SL) or head
length (HL); n = 25.
Coloration of preserved material. The colour pattern is basically the same as that of G. a n j e re n s i s (see that
species’ account), but differs from it in that the pectoral fin is plain dusky, with no rows of dark brown speckles; the
dusky anal fin may have distinct rounded to elongate dark brown spots along the centre of the fin, or a dark streak
present on the membrane parallel to the fin rays; and the blackish line on the dorsal surface of the eye is usually
split into two thin or diffuse blackish lines (often diverging), which may be indistinct or broken-up (Fig. 12D). The
dark brown mark (which may be indistinct) over the pectoral fin base is a blotch surrounding a small pale central
spot (spot often indiscernible in preserved specimens) or short brown bar running below pale central spot.
Distribution. This species is known only from the Hawaiian island chain. Thacker (2004) identified some
specimens from Rarotonga (Cook Islands) and Moorea (Society Islands) as G. k n i g h t i , based by their having a
“dash-shaped mark on the shoulder”. These specimens were used in the 2004 genetic analysis and appear as sister
to the Hawaiian specimens of G. k n i g h t i (Thacker 2004: Fig. 2; Thacker pers. comm.). However, we have seen no
specimens identifiable as G. k n i g h t i from these localities (only G. c a u e re n s i s ).
Ecology. Most specimens have been taken from shallow habitats, tidepools or “brackish pools”, at depths of
0–5 m, from sand and mangrove areas.
Smith (1989) compared the alarm response of Hawaiian Gnatholepis anjerensis (= G. k n i g h t i ) to that of
Asterropteryx semipunctatus, a sympatric gobiine, and found that Gnatholepis did not appear to have an alarm
Paratype
(male)
Means
(males)
Maximum
(males)
Minimum
(males)
Means
(females)
Maximum
(females)
Minimum
(females)
Standard length 43.0 45.9 64.0 32.5 47.0 61.0 34.0
Head length in SL 26.7 26.8 27.9 25.0 27.8 29.9 26.3
Head depth in HL 72.2 76.3 85.6 69.5 73.0 76.4 66.3
Head width in HL 67.8 72.0 78.3 67.4 70.0 80.0 61.2
Body depth at anus in SL 21.6 22.5 24.6 21.0 23.8 26.0 20.6
Body depth at D1 origin in SL 22.3 23.2 25.1 21.0 24.3 26.1 22.4
Caud. ped. length in SL 17.0 16.6 17.5 15.2 17.3 19.0 15.2
Caud. ped. depth in SL 12.1 11.9 13.1 10.2 11.8 13.5 10.3
Snout length in HL 33.0 35.5 43.2 29.9 37.1 44.0 28.6
Eye width in HL 25.2 27.9 32.0 25.0 27.3 30.6 24.8
Jaw length in HL 34.8 36.1 40.6 31.6 34.7 37.1 29.4
Interorbital in HL 9.6 9.0 10.4 7.2 8.2 9.7 6.1
Preorbital width in HL 19.1 21.9 24.5 19.1 23.1 27.9 20.4
Pectoral fin in SL 26.7 26.4 27.9 24.4 25.3 28.5 23.1
Pelvic fin in SL 28.4 27.1 29.6 24.7 24.9 28.2 22.6
Caudal fin in SL 32.3 31.3 35.6 27.8 28.3 30.1 26.1
Adpressed D1 in SL 24.9 23.3 25.9 14.7 23.3 25.6 21.1
3
rd
D1 fin spine in SL – 16.7 18.4 14.5 16.1 17.9 14.9
4
th
D1 fin spine in SL 16.3 17.8 18.9 16.0 16.6 18.8 14.7
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
pheromone (which Asterropteryx did). Cole (1990) described the sex ratio and gonad structure of this species (as G.
anjerensis; her material was all from Hawaii).
Comparisons. Gnatholepis knighti is very similar to G. a n j e re n s i s but has cycloid scales on the predorsal
region and the transverse black line on upper part of eye is often split in two (always a single line in G. a n j e r e n s i s )
(Fig. 12B, D).
The very brief description of G. k n i g h t i provided in Thacker (2004b) emphasises the presence of a dash-shaped
dark mark or line dorsal to the pectoral fin as being diagnostic for the species. From our examination of specimens
and colour photographs of live and freshly dead Gnatholepis, we found that this mark to be variably developed in
both G. a n j e r e ns i s and G. k n i g h t i .
Specimens from Rarotonga and Moorea identified by Thacker as G. k n i g h t i appeared as sister to the Hawaiian
G. k n i g h t i in a strict parsimony consensus tree (Thacker 2004a: Fig. 2; Thacker pers. comm.). Thacker et al.
presented further genetic analysis of a smaller number of Gnatholepis specimens in which the same groups
(Rarotonga/Moorea and Hawaii) were sisters in the parsimony consensus tree but the Rarotonga/Moorea
specimens were sister to G. a n j e r e ns i s in the Bayesian analysis (Thacker et al. 2008: fig. 1). As the specimens no
longer exist it is not possible to discover what these fish actually were.
Remarks. The holotype (USNM 50653) has been noted as missing since 1948 and may have never been
received by USNM. Thacker (2004b) remarked, but not Randall and Greenfield (2001) nor Greenfield and Randall
(2004), that the holotype of G. k n i g h t i is lost. However, 10 lots of paratypes exist in six American museums: CAS-
SU 7468, FMNH (5), ANSP 24222 (1, missing), MCZ 28903, USNM 144094 and AMNH 2294. The holotype was
illustrated by Jordan and Evermann (1905: 360, pl. 58).
The oldest material of G. k n i g h t i was collected in Oahu by the medical officers aboard the USS Portsmouth in
1873–74 (Jordan and Evermann 1905).
Gnatholepis ophthalmotaenia (Bleeker, 1854)
(Figs 12E, 21–23; Tables 5–8, 13)
Gobius ophthalmotaenia Bleeker, 1854: 46 (Nova Selma, Cocos-Keeling Island).
Gnatholepis davaoensis Seale, 1910: 537 (Samal Island, Davao Gulf, Mindanao).—Randall and Lim 2000: 638; Randall and
Greenfield 2001: 12 (Taiwan); Senou et al. 2004: 243 (Japan).
Gnatholepis gemmeus Herre, 1927: 135, pl. 9, fig. 3 (Samal Island, Davao Gulf, Mindanao).—Herre 1931: 14 (New Hebrides);
Herre 1935: 357–358 (Bushman Bay, Malekula Island, New Hebrides); Koumans 1940: 184.
Gnatholepis corlettei Herre, 1935: 418 (Bushman’s Bay, Malekula, New Hebrides).—Herre 1931: 14 (nomen nudum); Herre
1936: 356–357 (Bushman Bay, Malekula Island, New Hebrides); Koumans 1940: 138; Böhlke 1953: 112; Ibarra and
Stewart 1987: 40.
Gnatholepis deltoides—Masuda et al. 1984: 252, plate 240Q (Yaeyama Islands).
Gnatholepis anjerensis (in part)—Randall and Greenfield 2001: 3.
Gnatholepis cf anjerensis—Kuiter and Tonozuka 2001: 671 (S of Bitung, Sulawesi).
Gnatholepis ophthalmotaenia—Kuiter and Tonozuka 2001: 672, figs A–C (Bali, Indonesia); Allen and Adrim 2003: 58 (Flores,
Ambon, Halmahera).
Gnatholepis anjerensis—Motomura et al. 2010: 213, fig. 524 (Yaku-shima Island).
Diagnosis. A moderate-sized (up to 55 mm SL) Gnatholepis with distinctive colour pattern in males of six broad
vertical dark blotchy bars along mid-side of body, several staggered rows of black spots along the dorsal part of the
body, one to four irregular rows of golden yellow (in life) spots along lower half of body, two characteristic small
black spots at base of first dorsal fin and one to three staggered rows of oval black ocellate spots on anal fin (in life,
red, blue and yellow may surround black spots), females with vertical dark body bars much paler and red and
yellow markings much less distinct; second dorsal and anal fin rays nearly always I,11; pectoral rays 15–17,
usually 16; lateral scales 24–30, usually 26; 9–10 predorsal scales (usually 8–9), ctenoid and cycloid.
Material Examined. COCOS-KEELING: Syntype of Gobius ophthalmotaenia, RMNH 4526, 55 mm SL
male [one of 5 possible syntypes], Nova-Selma, J. Clunies-Ross. JAPAN: NSMT-P67349, 1(33), Hoshizuma-no-
hama Beach, Iriomote-jima, Yaeyama Islands, T. Mukai, 17 August 2000; NSMT-P64595, 3(22.5–29), Beguala
Bay, Ambon Island, 1 December 1988; NSMT-P64639, 1(27), Kotania Bay, Seram Island, 3 December 1988;
NSMT-P64505, 1(25.5), SE coast Kotania Bay, Seram Island, 4 December 1988. TAIWAN: Neotype of
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Gnatholepis davaoensis, 52 mm SL male, BPBM 18670, tidepool at S end of island at Hou Pi Hoo, J. Randall, A.
Choi and C. Wong, 11 June 1975; ASIZP 59337, 3(39–46), Wan-li-tung, Kenting National Park, Henchuen,
Pingtung County, K.T. Shao, 8 October 1985. BPBM 38374, 4(27–52), same data as neotype. PHILIPPINES: AMS
I.21938-003, 16(10.5–35.5), inlet 2 km S of Tambuli, Mactan Island, Cebu, D. Hoese and T. Hynes, 2 May 1980;
AMS I.24136-001, 1(36.5), Batangas, Anilao, E. Murdy, 1 February 1980; AMS I.21931-004, 4(34–40), Lapu
Lapu market, Cebu, Mactan Island, E. Murdy, 30 April 1980; AMS I.21931-007, 1(36), Lapu Lapu market, Cebu,
Mactan Island, E. Murdy, 30 April 1980; ex-AMS I.21931-005, 2(38–42), same data as previous. INDONESIA:
NSMT-P64545, 3(22.5–29), Baguala Bay, Ambon, K. Shibukawa, 1 December 1998; USNM 266393, 1(35.5),
Kampung Pasir Putih, Jailolo district, Halmahera Island, P. Taylor, 30 April 1981; NSMT-P64505, 2(17–26.5), SE
coast of Kotania Bay, Ceram, K. Shibukawa, 4 December 1998; NSMT-P64639, 1(27), Kotania Bay, Ceram, local
fisherman, K. Matsuura and K. Shibukawa, 3 December 1998; NTM S.12697-001, 1(29), small creek at beach at
Poso Harbour, Sulawesi, H. Larson and R. Williams, 5 September 1989. PAPUA NEW GUINEA: AMS I.32492-
004, 1(29.5), mangrove at Madang, 29 November 1987; AMS I.32492-009, 1(26), mangrove at Madang, 29
November 1987; NTM S.13679-031, 1(33.5), opposite Christensen Research Institute, Nagada Harbour, Madang,
H. Larson and M. Jebb, 17 October 1992; USNM 308210, 21(11.5–34), Maiwara Island, W of Alotau, Milne Bay,
T. Roberts, 3 September 1975. SOLOMON ISLANDS: AMS I.31088-002, 1(25), Lembu, New Georgia, S. Blaber,
August 1988; CAS 51506, 23(29–35.5), first point S of Paeu River, Vanikolo, Santa Cruz Islands, R. Bolin, 30
September 1958; NTM S.11328-002, 1(34), Talisondo Island, Vona Vona, S. Blaber, 9 July 1987. VANUATU:
FMNH 17367, holotype of Gnatholepis corletti, 25 mm SL female, Bushman’s Bay, Malekula Island, New
Hebrides, A.W. Herre, 4 April 1929.
Other material; no data taken. PHILIPPINES: USNM 160804, 34; CAS 51508, 8, Dumaguete Beach,
Negros Oriental; USNM 261647, 29, Negros Island, Negros Oriental. INDONESIA: USNM 243456, 2, Ambon,
Moluccas.
Description. Based on 54 specimens, 19–55 mm SL.
First dorsal VI; second dorsal I,9–I,11 (mean I,10.9); anal I,10–11 (mean I,10.9), pectoral rays 15–17 (mean
16.0), segmented caudal rays nearly always 17; caudal ray pattern nearly always 9/8; branched caudal rays 6/6 to 8/
6 (usually 7/6); lateral scale count 24–30 (mean 26.4); TRB 9–11 (mean 9.8); predorsal scales 6–10 (mean 8.5);
circumpeduncular scales 11–12 (mean 12.0). Gill rakers on outer face of first arch 1–2 + 3–5 (in 7, usually 1+4).
Body compressed, width at anus 10.9–24.4% (mean 14.0%) of SL. Body rather stocky, body depth at anus
20.9–26.9% (mean 23.9%) of SL, body depth at first dorsal fin origin 11.2–27.5% (mean 24.1%) of SL. Head
compressed, broader ventrally, slightly deeper than wide, HL 25.2–30.7% (mean 28.0%) of SL; head depth at
posterior preopercular margin 68.5–86.1% (mean 75.8%) of HL; head width at posterior preopercular margin
57.7–78.3% (mean 69.2%) of HL; head profile bluntly pointed to somewhat rounded; nape profile slightly curved.
Mouth just terminal, fleshy snout tip slightly overhanging jaw tips, mouth slightly oblique; jaws generally reaching
to below anterior margin of eye; upper jaw length 28.1–42.0% (mean 34.4%) of HL. Upper lip smooth, narrower
than lower, lower lip finely papillose close to teeth, with twist or fold posteriorly, forming triangular flap, lip
interrupted at chin. Eye moderate, dorsolateral, 25.0–33.9% (mean 28.8%) of HL; preorbital width 16.5–24.0%
(mean 19.8%) of HL. Snout bluntly pointed, 25.3–38.3% (mean 32.2%) of HL; posterior naris round to almost
triangular, adjacent to anterior margin of eye; anterior naris in short tube, higher on posterior margin of eye, at level
with lower half of eye. Interorbital narrow, 6.3–11.0% (mean 8.1%) of HL. Caudal peduncle compressed, length
14.6–19.4% (mean 16.7%) of SL; caudal peduncle depth 10.9–14.0% (mean 12.1%) of SL.
First dorsal fin rounded to square, with no spines greatly elongate; first or second spine usually longest; when
adpressed, spine tips reaching to first to second element of second dorsal fin. First dorsal spine length 15.5–21.8%
(mean 19.2%) of SL; second dorsal spine length 15.9–22.2% (mean 18.6%) of SL; third dorsal spine length
14.5–22.7% (mean 17.8%) of SL. Second dorsal fin as tall as first dorsal fin, rays longer posteriorly, fin pointed
posteriorly. Anal fin a little lower than second dorsal fin, anteriormost rays shorter than posterior rays; fin pointed
posteriorly. Second dorsal and anal fin rays, when adpressed, usually reaching caudal fin in adults. Pectoral fin
somewhat pointed, central rays longest, 23.3–29.0% (mean 26.4%) of SL; fin reaching back to above first few anal
fin rays. Pelvic fins fused, frenum with distinctive finely fimbriate margin, fins oval, reaching back to anal fin
spine, 22.2–30.0% (mean 27.4%) in SL. Caudal fin oval to rounded, 27.9–35.0% (mean 31.4%) of SL.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
FIGURE 21. Gnatholepis ophthalmotaenia male, Moalboal, Cebu, Philippines. Photo by Robert Patzner.
Gill opening restricted, extending anteriorly to lower edge of pectoral base. Gill rakers as in G. a n j e re n s i s .
About one-quarter of first gill arch bound by membrane to inner face of opercle. Tongue short, tip bilobed.
Teeth in upper jaw in two to three rows across front and one row at side of jaw, outermost row teeth largest,
curved (less so at side of jaw) and pointed, largest teeth at front of jaw; inner row teeth very small, sharp and evenly
sized. Teeth in lower jaw in two to three rows, arranged similarly to upper jaw; posteriormost one or two outer row
teeth enlarged, often recurved in males.
Ctenoid scales on predorsal area of variable extent, from just over opercle to close up to behind eyes, midline
often of cycloid scales; occasionally entire predorsal covered with cycloid scales. Opercle covered with scales,
usually mostly ctenoid. Preopercular scales usually cycloid, although upper or rear part of preopercle may include
ctenoid scales; cycloid scales often extending anterior to vertical dark cheek-bar below eye. Breast with cycloid
scales usually reaching forward to anterior of preopercular margin (below eye); scales reaching at least to below
rear margin of preopercle. Pectoral fin base covered with cycloid scales. Belly scales along midline cycloid
anteriorly at least; posterior half of belly midline may be of ctenoid scales.
Head pores and sensory papillae as in G. a n j e re n s i s (Fig. 1).
Coloration of fresh material. Illustrations of living fish can be found in Senou et al. (2004: 243, as G.
davaoensis), Kuiter and Tonozuka (2001: 672, figs A–C) and Randall and Greenfield (2001: 9F, G, as G.
davaoensis).
Head and body bright pearly white to yellowish white with six broad diffuse black to brown bars or rounded
blotches along mid-side of body, reducing in size posteriorly, posteriormost small blotch at mid-base of caudal
peduncle; often with five indistinct brown short saddles (or staggered pairs of rectangular blotches) crossing
dorsum (Fig. 21). Three or more irregular rows of small round black spots along dorsal third of body, often with
additional two to four rows of round dull yellow to orange spots along remainder of side of body (often a round
orange spot placed in centre of each black mid-lateral bars or blotches).
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FIGURE 22. Gnatholepis ophthalmotaenia female, Madang, New Guinea. Photo by Gerry Allen.
Predorsal and snout finely speckled with dark brown to black, opercle and cheek variably spotted with bright
blue-white to pearly white spots that extend onto body, forming several rows of blue spots, these spots often
arranged around each mid-lateral black blotch like an ocellus. Dark brown to black cheek-bar of variable width,
from ventral edge of eye running vertically to slightly obliquely down to end on lower preopercular edge, but not
extending onto branchiostegal membranes; sometimes black cheek-bar narrow and surrounded by diffuse brown to
red-brown pigment. Black to very dark brown line from above middle of iris running obliquely back over top of
eye. Above pectoral fin base, indistinct and diffuse brown to dull orange blotch present, partly connected to broken
brown line or series of brown spots running from behind eye along top of preopercle. Pectoral fin base white with
pale blue-bordered horizontal brown line or diffuse orange to light brown blotch across mid-base.
First dorsal fin translucent, with three to six rows of short oblique dusky streaks or spots, beginning of first
three (or more) wavy rows marked by black spot on front of first spine and often one black spot at base of spine;
two dense round black spots at base of fin, one at base of fourth spine and one at base of sixth. Second dorsal fin
similar but with more rows of diffuse dark spots which may transform into darker spots and vertical streaks on
membrane, especially on posterior half of fin; beginning of rows of spots marked by black spot on front of second
dorsal fin spine. Anal fin brightly coloured in male: dull white to blue, with one to three staggered black ocellate
oval to round spots, becoming larger and more numerous toward rear of fin; black spots surrounded by red to pink
and interspersed with pale blue to yellow round to oval spots (female less colourful, with less blue and yellow
spotting on all parts of body and fins) (Figs 21–23). Caudal fin translucent with small red-brown blotch on mid-
base, and irregular rows of small black spots or diffuse light brown blotches. Pectoral fins transparent with white
fin rays and fine white speckling on basal part of fin. Pelvic fins white to yellow, with pale bluish to white margin.
A photograph by Gerry Allen of a female from Madang, New Guinea, shows a very plain fish that looks like G.
anjerensis but for the round orange and pale blue spots on its side, the dark spots on the anal fin and the two
diagnostic black spots at the base of its first dorsal fin (Fig. 22).
Coloration of preserved material. Head and body yellowish white with five somewhat indistinct brown
saddles (may be split into pairs) crossing upper third of body, six vertical broad diffuse brown bars or large blotches
along mid-side of body, posteriormost bar or blotch at mid-base of caudal peduncle. Three irregular rows of small
round black to dark brown spots along dorsal third of body, some specimens with additional four rows of round
brown spots along remainder of side of body (these spots never as dark as those in uppermost three rows).
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Predorsal and snout finely speckled and spotted with brown, opercle and cheek mottled with dusky pigment.
Narrow blackish cheek bar from ventral edge of eye running vertically to slightly obliquely down to end on lower
preopercular edge, but not extending onto branchiostegal membranes; narrow stripe usually surrounded by diffuse
brownish area. Blackish to dark brown line from above middle of iris running obliquely back over top of eye, may
meet its counterpart in interorbital space. Above pectoral fin base, diffuse brownish blotch may be present, partly
connected to brown line or series of brown spots running from behind eye along top of preopercle. Pectoral fin base
with wavy to diffuse horizontal brown line across dorsal half, line extending onto lower part of fin and ending in
small brownish blotch.
First dorsal fin transparent, with three to four wavy brown lines or series of spots, beginning at first spine;
beginning of first three wavy lines marked by characteristic blackish spot on front of spine; two diagnostic dense
black spots at base of fin, one at base of fourth spine and one at base of sixth. Second dorsal fin similar but with
four to five rows of dark brown to blackish spots which may transform into series of darker spots and vertical
streaks on membrane, especially on posterior half of fin, blackish spots becoming more intense posteriorly and
proximally. Anal fin dusky, with one to three staggered rows (more rows in larger fish) of black ocellate oval spots,
becoming larger and more numerous toward rear of fin. Caudal fin translucent with small brown blotch on centre of
basal scale sheath, and four to five evenly spaced blackish oval spots forming vertical to curved line across fin ray
bases; fin covered with irregular rows of small blackish to brown spots and streaks. Pectoral fins transparent to
translucent dusky, may be finely mottled with diffuse brown in larger specimens. Pelvic fins plain dusky to almost
black, may have white margin.
Distribution. Gnatholepis ophthalmotaenia has a relatively restricted distribution, being known from the Cocos-
Keeling Islands, Philippines, Taiwan, southern Japan, Indonesia, New Guinea, Solomon Islands and Vanuatu.
FIGURE 23. Gnatholepis ophthalmotaenia female, Moalboal, Cebu, Philippines. Photo by Robert Patzner.
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Ecology. This species does not seem to be as common or abundant as G. a n j e r en s i s or G. c a u e re n s i s . It is found
in shallow waters, from reef tidepools, among seagrass, in shallow mangroves and (usually) sand and coral rubble
reef substrates at depths of 0.2–12 m.
Comparisons. This species is easy to distinguish from all other Gnatholepis with scaled cheeks and opercles,
due to the irregular rows of ocellate black spots on the anal fin (there is no equivalent present in other species), the
nape scales being mostly ctenoid and the breast scales extending forward of the rear edge of the preopercle. It is a
relatively small to moderate-sized species also (largest only 55 mm SL) and is sexually dichromatic (unlike
Gnatholepis species other than G. a r g u s ), with females showing much reduced patterning and intensity of colour
compared to males.
TABLE 13. Measurements of specimens of Gnatholepis ophthalmotaenia, expressed as percentage of standard length
(SL) or head length (HL); n = 53.
Remarks. The syntypes of Gnatholepis ophthalmotaenia Bleeker are presently included in RMNH 4526,
which in 1988 contained 13 specimens (29–49.5 SL), five of which are within Bleeker’s size range of 58–64 TL
(he based his description on four specimens, 58–64 mm TL). The senior author’s notes state that all specimens had
colouring well-preserved and that the colouring was similar to that shown and annotated for G. o p h t h a l m o t a e n i a in
Bleeker (1983), with the “very fine black dots on upper head and back” to be added by the artist. Randall and
Greenfield (2001) did not provide the basis of their synonymy of G. o p h t h a l m o t a e n i a with G. a n j e r e n s i s and type
material was apparently not examined.
The holotype of Gnatholepis davaoensis Seale, 1910, from Samal Island, Mindanao, was listed as lost (“…. is
no longer in the Bureau of Science collection”) by Herre (1927). A neotype (BPBM 18670) from Taiwan was
designated by Randall and Greenfield (2001), as they considered that G. d a v a o e n s i s was the correct name to apply
to G. o ph t h a l m o t a e n i a .
The holotype and all 24 paratypes of Gnatholepis gemmeus, Herre, 1927, were destroyed in WWII. Koumans (1940)
saw three bottles at the Manila Bureau of Science, all marked as types (from Samal, Davao, Dumaguete and Sitankai).
Means
(males)
Maximum
(males)
Minimum
(males)
Means
(females)
Maximum
(females)
Minimum
(females)
Standard length 34.6 55.0 22.5 26.8 35.5 19.0
Head length in SL 27.5 30.7 25.2 28.8 30.0 27.4
Head depth in HL 76.9 86.1 68.5 74.3 82.3 68.7
Head width in HL 69.1 78.3 57.7 69.3 76.0 57.8
Body depth at anus in SL 24.2 26.9 22.2 23.5 25.3 20.9
Body depth at D1 origin in SL 23.4 26.0 11.2 25.0 27.5 12.2
Caud. ped. length in SL 16.8 19.2 14.7 16.5 19.4 14.6
Caud. ped. depth in SL 12.3 14.0 11.2 11.8 13.0 10.9
Snout length in HL 32.7 38.3 25.3 31.1 37.0 26.9
Eye width in HL 28.2 31.5 25.0 29.6 33.9 26.0
Jaw length in HL 35.4 42.0 29.0 32.4 37.3 28.1
Interorbital in HL 8.3 11.0 6.3 7.9 10.1 6.3
Preorbital width in HL 19.6 22.1 16.5 20.0 24.0 17.4
Pectoral fin in SL 26.3 29.0 23.3 26.5 28.9 24.2
Pelvic fin in SL 27.4 30.0 22.2 27.2 29.3 23.4
Caudal fin in SL 31.9 35.0 27.9 30.1 31.9 28.2
Adpressed D1 in SL 23.7 26.9 21.4 21.8 23.7 18.5
3
rd
D1 fin spine in SL 18.3 22.7 15.8 16.8 18.7 14.5
4
th
D1 fin spine in SL 17.4 20.7 14.6 15.7 17.4 13.6
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
The specimens identified as G. a n j e r e n s i s from Yaku-shima Island, Japan, in Motomura et al. (2010) have not
been examined. However, the record is placed here based on Motomura et al.’s (rather dark) Figure 524. This fish
has the characteristic anal fin red and black ocellate markings of G. o p h t h a l m o t a e n i a and the two black spots along
the base of the first dorsal are just visible.
Gnatholepis pascuensis Randall and Greenfield, 2001
(Fig. 24; Tables 5–8)
Gnatholepis sp. nov.—DiSalvo et al. 1988: 461 (Easter Island).
Gnatholepis cauerensis pascuensis Randall and Greenfield, 2001: 11–12, pl. IIE (Easter Island, off Tahai).—Randall et al.
2005: 47.
Gnatholepis pascuensis—Randall 2009: 179.
Diagnosis. A rather small Gnatholepis (up to 45 mm SL) with distinctive colour pattern of two rows of five
elongate dense black blotches on side of body, anteriormost three blotches largest, joined by faint broken yellowish
brown lines in life, very broad black cheek-bar and broad black eye-stripe (both as wide as pupil) crossing
interorbital space; ctenoid body scales reaching rear corner of opercle, or to above pectoral fin base; nape midline
scales always cycloid; few ctenoid scales present on opercle; distinct flap present at end of lower lip; 18–19
pectoral fin rays.
FIGURE 24. Gnatholepis pascuensis, Easter Island. Photo by Jack Randall.
Material examined. EASTER ISLAND: Holotype BPBM 32850, 34 mm SL, off Tahai, J. Randall and party,
13 February 1985; BPBM 6743, 1(30), offshore from Akapu, J. Randall and B. Baker, 3 February 1969; BPBM
32852, 1(41.5), off Tahai, J. Randall and party, 15 February 1985; BPBM 32853, 1(45), between Motu Tautara and
Ara O Hara, J. Randall and party, 15 February 1985; BPBM 32851, 2(35–39), off E end Anakena, J. Randall and
party, 14 February 1985.
Remarks. Randall and Greenfield (2001) created the name Gnatholepis cauerensis pascuensis based on one
specimen (BPBM 32850). Randall (2009) subsequently elevated G. p a s c u e n s i s to full species status and presented
two additional lots as paratypes (BPBM 6743 and BPBM 32851–53). These specimens were not so designated in
the original description (were given as “Other Material”) and therefore do not hold any type status. It is also
unfortunate that Randall and Greenfield chose a specimen in poor condition for the holotype when there were three
better specimens available (e.g. BPBM 32853, 32851).
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We agree with Randall (2009) in assigning full species status to G. p a s c u e n s i s . It has a high pectoral fin ray
count (18–19). Although Randall (2009) reported one specimen with 17 rays on one side, the six specimens that we
examined had 18–19 rays on both sides. Gnatholepis pascuensis also has a characteristic intense colour pattern of
two rows of five elongate black blotches laterally, a very broad black cheek-bar and broad black eye-bar (nearly as
wide as pupil) crossing the interorbital space (Fig. 24). Some G. c a u e re n s i s specimens also have a broad cheek and
eye-bar but have six lateral dark blotches and lower pectoral fin ray counts.
Gnatholepis cauerensis specimens from Rapa and Pitcairn have also 17–19 pectoral rays but are typically pale
in colour and do not show any significant colour differences from other G. c a u e re n s i s .
Gnatholepis thompsoni Jordan, 1904
(Figs 4E–F, 12F, 25–26; Tables 5–8, 14)
Gnatholepis thompsoni Jordan, 1904: 541, pl. 1, fig. 2 (Bush Key, Tortugas Archipelago, Florida).—Böhlke 1953: 113; Randall
1968: 248–249 (Bermuda; Caribbean Sea); Lubbock 1980: 296 (Ascension Island); Robins and Ray 1986: 246 (Bahamas);
Edwards and Glass 1987: 659 (St Helena); Miller 1990: 931; Smith-Vaniz et al. 1999: 316 (Bermuda); Schwartz 1999: 284
(North Carolina); Brito and Miller 2001: 255, Fig. 2A (Cape Verdes); Gasparini and Floeter 2001: 1646 (Trindade Island,
Brazil); Rocha and Rosa 2001: 992 (Maranhão, Brazil); Randall and Greenfield 2001: 1; Thacker and Cole 2002: 840;
Araújo and Freitas 2002: 2 (Madeira Island); Collette et al. 2003: 122 (Navassa Island); Murdy and Hoese 2003: 1795;
Smith et al. 2003: 62 (Belize); Ross and Rhode 2004: 309, plate IIId (Onslow Bay, North Carolina); Nelson et al. 2004:
171; Rocha et al. 2005: 1–6.
Gnatholepis sp.—Lubbock in Miller 1990: 931 (Ghana).
Bathygobius soporator—Debelius 1997: 244 (Cape Verde Islands) [misidentification].
Gnatholepis scapulostigma—Brito and Miller 2001: 258 (Cape Verde Islands).
Diagnosis. A moderately large Gnatholepis (up to 58 mm SL) with nape midline scales always cycloid and most of
predorsal scales cycloid; body pale with 6–8 rows of staggered dark brown spots, mid-lateral spots may be largest;
transverse black line on the upper part of the eye joining somewhat oblique to curved black line or bar crossing
cheek and ending well behind end of jaw; third to fourth first dorsal fin spines longest, fin with square to
rectangular appearance when extended; second dorsal and anal fin rays usually I,11; pectoral rays 16–18, usually
17; lateral scales 26–29, usually 27; 9–11 predorsal scales (usually 10), all cycloid.
Material examined. BELIZE: USNM 276135, 12(10–41.5), E side Carrie Bow Cay, D.J. Johnson and P.
Keener, 12 November 1984; USNM 276131, 4(18.5–30.5), S end Carrie Bow Cay, D.J. Johnson and P. Keener, 5
November 1984; USNM 346472, 2(27–33.5), Carrie Bow Cay, D. Smith, C. Thacker, 16 September 1997.
NICARAGUA: USNM 362225, 3(24.5–30), Narrow Cay, Serrana Bank, Oregon Cruise 78, 20 May 1962; USNM
320763, 2(27–36.5), off Corn Island, Caribbean, Oregon Cruise 78, 2 June 1962. BAHAMAS: USNM 198783,
1(33.5), S end of beach near canal inlet, Lyford Cay, Clifton Bay, W.L. Schmitt, 16 August 1961. CUBA: USNM
82520, 6(29–44), San Antonio, Tomas Barreras Expedition, Henderson and Bartsch, 4 June 1914. JAMAICA:
LACM 5974, 12(34.5–58), Pedro Cays, D.K. Caldwell, 21 April 1959. LESSER ANTILLES: USNM 264886,
1(58), E side Cocoa Point, Barbuda, 25 April 1959. VENZUELA: USNM 179253, 1(41.5), Sarqui Islands, Los
Roques Islands, P. Bottome and Wallis, 4 July 1958; USNM 179252, 11(31.5–51), Yonqui, Los Roques Islands, P.
Bottome and Wallis, 5 July 1958; USNM 179254, 3(14–37), Sarky Island, Los Roques, P. Bottome and Wallis, 27
June 1958. TRINIDAD AND TOBAGO: USNM 317097, 10(11–41), leeward side Little Tobago Island, J.T.
Williams and party, 6 September 1990; USNM 317098, 8(12–34), Man-of-War Bay, Booby Island, Tobago, J.T.
Williams and party, 14 September 1990. BRAZIL: USNM 357711, 3(34.5–40.5), Atol das Rocas, Rio Grande do
Norte, N.A. Menezes, February 1972; USNM 368819, 1(28), Bay of Turiacu, 9 March 1963. ASCENSION
ISLAND: USNM 368871, 1(30), rocky point at N edge of English Bay, R. Manning, P. Kashulines, 20 May 1971;
USNM 218840, 1(44), McArthur Point, M.L. Jones and party, 11 July 1976. ST HELENA: USNM 267896,
6(25–36), Lemon Valley Bay, A. Edwards and C. Glass, 28 June 1983.
Other material; no data taken. ASCENSION ISLAND: USNM 368859, 2.
Description. Based on 61 specimens, 21.5–58.0 mm SL. An asterisk indicates the counts of the holotype of
Gnatholepis thompsoni (data provided by Dave Catania of CAS).
First dorsal VI*; second dorsal I,10–I,12* (mean I,11.0); anal I,10–12 (mean I,11.0*), pectoral rays 16*–18
(mean 16.9), segmented caudal rays always 17; caudal ray pattern nearly always 9/8; branched caudal rays 7/6 to 8/
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
7 (usually 7/6); lateral scale count 26–29 (mean 27.0*); TRB 9*–12 (mean 10.4); predorsal scales 9–11.5 (mean
9.8, 10 in holotype); circumpeduncular scales 12–14 (mean 12.1). Gill rakers on outer face of first arch 1–2 + 3–5
(in 8, usually 1+4).
FIGURE 25. Gnatholepis thompsoni, Florida, USA. Photo by Rob Myers.
Body compressed, width at anus 9.8–16.0% (mean 13.2%) of SL. Body rather slender, body depth at anus
17.7–25.7% (mean 21.5%) of SL, body depth at first dorsal fin origin 19.3–25.3% (mean 22.4%) of SL. Head
compressed, slightly broader ventrally, deeper than wide, HL 25.7–30.0% (mean 28.0%) of SL; head depth at
posterior preopercular margin 58.5–76.6% (mean 67.5%) of HL; head width at posterior preopercular margin
57.3–77.2% (mean 67.7%) of HL; head profile bluntly pointed to rounded; nape profile slightly curved to flat.
Mouth subterminal to nearly terminal, slightly oblique; jaws generally reaching to below anterior half of eye; upper
jaw length 28.1–39.0% (mean 34.4%) of HL. Upper lip smooth, narrower than lower, lower lip with papillose ridge
close to teeth, with twist or fold posteriorly, forming triangular flap, lip narrowly interrupted at chin. Eye moderate
to relatively small, dorsolateral, 23.0–35.5% (mean 27.0%) of HL; preorbital width 13.6–24.8% (mean 19.4%) of
HL. Snout bluntly pointed to rounded, 25.7–38.2% (mean 30.9%) of HL; posterior naris rounded, close to anterior
margin at middle of eye; anterior naris in short tube, higher on posterior margin of eye, about level with middle of
eye or somewhat ventral to it. Interorbital narrow, 4.9–11.0% (mean 7.9%) of HL. Caudal peduncle compressed,
length 14.3–18.6% (mean 16.5%) of SL; caudal peduncle depth 10.2–13.8% (mean 11.6%) of SL.
First dorsal fin rounded to roughly triangular, nearly square in large adults, with no spines elongate; second to
fourth spine longest; when adpressed, spine tips reaching to first to third element of second dorsal fin. Second
dorsal spine length 14.9–20.3% (mean 17.2%) of SL; third dorsal spine length 15.8–21.6% (mean 18.0%) of SL.
Second dorsal fin as tall or nearly as tall as first dorsal fin, posteriormost rays usually longer than anterior, fin
pointed to slightly rounded posteriorly. Anal fin lower than second dorsal fin, anteriormost rays shorter than
posterior few rays; fin usually pointed posteriorly. Second dorsal and anal fin rays, when adpressed, just reaching
caudal fin rays. Pectoral fin oval to somewhat pointed, central rays longest, 22.5–29.8% (mean 26.0%) of SL; fin
reaching back to above first few anal fin elements. Pelvic fins fused, frenum with distinctive finely fimbriate
margin, fins round to somewhat oval, reaching to first one or two anal fin elements, 23.3–31.0% (mean 27.2%) in
SL. Caudal fin moderate, oval, rounded to slightly pointed, 26.6–37.4% (mean 31.8%) of SL.
Gill opening restricted, extending anteriorly to lower edge of pectoral base or to just under opercle. One or two
slender, fleshy gill rakers on outer face of first arch, close to angle of arch, remaining few rakers shorter, pointed;
outer rakers on second gill arch consisting of two series of papillae, one low and one (mostly anterior) of slightly
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pointed papillae; outer rakers on remaining arches low, stubby, broad-based. Inner face of upper limb of first gill
arch, and to lesser extent, upper limbs of other arches, covered with low dense fleshy papillae, usually forming
clumps or groups; dorsal portion of first arch may have short fleshy protuberances ending in one or several
papillae; inner face of second arch smooth, without papillae but with about 12 short fleshy rakers with rounded,
finely papillose tips. About one quarter or less of first gill arch bound by membrane to inner face of opercle.
Tongue short, tip bilobed to concave.
Teeth in upper jaw in two to three rows across front and one row at side of jaw, outermost row teeth largest,
curved and pointed, largest teeth at front of jaw on either side of symphysis (smaller in females); innermost row
teeth very small, sharp and evenly sized. Teeth in lower jaw in two to three rows, arranged similarly to upper jaw
but outer row teeth may be smaller (in females); posteriormost one or two outer row teeth enlarged and somewhat
recurved in males.
Predorsal scales cycloid, ctenoid body scales reaching forward to rear corner of opercle to just over opercle, in
two specimens (from Brazil and Ascension), ctenoid scales reaching forward to over preopercle; nape midline
scales always cycloid (Fig. 4E–F). Opercle covered with cycloid scales, a few ctenoid scales on upper part may be
present (Jamaica specimens with mostly ctenoid scales). Preopercular scales cycloid, occasionally extending
anterior to vertical dark cheek bar below eye. Breast with cycloid scales reaching up to below mid-opercle or to
below rear half of opercle. Pectoral fin base covered with cycloid scales; occasionally with few ctenoid scales in
centre. Belly scales along midline usually cycloid; ctenoid scales may be present posteriorly.
Head pores as in G. a n j e re n s i s (Fig. 1).
Sensory papillae as in G. a n j e r e n s i s .
Coloration of fresh material. Published photographs of living or freshly dead fish can be seen in Debelius
(1997: 244, as Bathygobius soporator), Collette et al. (2003: 123, fig. 143) and Ross and Rohde (2004). Living and
freshly dead fish are very similar in colour to the spotted form of G. c a u e r e n s i s .
Description from colour photographs of living fish by Rob Myers (Figs 25–26). Head and body dull to pearly
white with six to eight rows of many small brown to red-brown round to vertically oval spots running length of
body, mid-lateral spots may be partly slightly larger and red-brown (may partly coalesce forming lines); rows of
spots underlain by five to six vertically oval, indistinct pale brown to purplish brown bars similar to those in G.
anjerensis. Predorsal lightly spotted with pale brown to red-brown. Side of head may be crossed horizontally by
red-brown or diffuse brown mottled line extending from just above rictus on to opercle; more often, opercle with
(often indistinct) brown to red-brown line from centre of preopercular margin horizontally or obliquely back to
opercle rear margin. Narrow to broad black to dark brown cheek bar from ventral edge of eye running vertically
down to end on lower preopercular edge. Cheek with diffuse brownish horizontal line crossing vertical cheek bar
(diffuse line may continue onto opercle as described above). Dense black to dark brown bar from middle to rear
half of iris running over top of eye. Above pectoral fin base, large black U-shaped flat-based blotch surrounding
round yellow spot; black blotch may be partly connected to thin red-brown line from behind eye along top of
preopercle; indistinguishable in small specimens. Pectoral fin base with horizontal red-brown line crossing middle
of fin base.
First dorsal fin translucent pale white, with four to seven rows of many pale purple-brown spots and short
streaks (usually forming short lines proximally. Second dorsal fin with similar five to seven rows of brown to
purple-brown spots. Anal fin yellowish white to white with blue-white margin; large adult from Florida with short
red-brown spot and short streaks along base of fin (difficult to see anal fin in most photos). Caudal fin translucent
white, with many irregular rows of small dark brown spots along fin membranes; posterior and ventral margins of
fin less pigmented. Pectoral fins transparent with translucent white rays and small fine white speckles basally.
Pelvic fins translucent white.
Coloration of preserved material. Based on typically coloured specimens from Jamaica (variation discussed
below). Head and body yellowish white to whitish with six or seven rows of many small brown spots running along
length of body, joined by variably developed faint brownish lines; rows or lines of spots faintly underlain by five to
six vertically oval, brownish bars similar to those in G. a n j e re n s i s ; in some specimens, second and third (counting
from dorsalmost) and fifth and sixth rows of spots joined by slightly darker background pigment, making these two
pairs of rows slightly more conspicuous. Dorsum crossed by about 9–10 usually very faint dusky square saddles
(first saddle on nape before first dorsal fin); these saddles not always discernible.
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
FIGURE 26. Gnatholepis thompsoni, San Salvador, Bahamas. Photo by Rob Myers.
Predorsal, snout and side of head faintly blotched and spotted with pale brown, small spots darkest. Opercle
with (often indistinct) pale brown line from centre of preopercular margin obliquely back to upper half of opercle
rear margin; line may curve up to rear corner of opercle. Narrow to broad black to light brown cheek bar from
ventral edge of eye running vertically to slightly obliquely down to end on lower preopercular edge, but not
extending onto branchiostegal membranes. Dense blackish to dark brown line from middle to rear half of iris
running over top of eye, often meeting its counterpart in interorbital space (Fig. 12F). Upper lip whitish, crossed by
short brownish bar (which may be joined by diffuse brown line starting from below anterior naris). Cheek with
diffuse brownish horizontal line or blotch crossing vertical cheek bar; brownish line variable in shape and intensity.
Above pectoral fin base, large brownish to dark brown, roughly U- or W-shaped flat-based blotch surrounding
small very pale brown round spot; brown blotch often partly connected to thin brown line (may be partly broken-up
or indistinct) running from behind eye along top of preopercle; pectoral blotch usually paler in small specimens.
Pectoral fin base with horizontal brown line crossing just above middle of fin base, line usually extending onto
lower part of fin.
First dorsal fin transparent to whitish, with four to six rows of light brown spots and short streaks (often
forming lines proximally), rows of spots beginning on first spine (darkest spots); lowermost one or two rows
darkest, especially anteriorly; distal part of fin may be plain whitish or translucent. Second dorsal fin with similar
five to seven rows of light brown spots or short horizontal streaks. Anal fin plain dusky to brownish, may be darker
distally. Caudal fin transparent to translucent brownish, sometimes with pale brown streaks or rows of fine brown
spots along fin membranes, especially near fin base and on upper half of fin. Pectoral fins transparent to translucent
dusky. Pelvic fins and frenum translucent plain pale brownish, membrane between rays may be darker than frenum.
Variation . Specimens from St Helena and Tobago are similar to the Jamaica fish but the background colour of
the head and body is brownish (not whitish or yellowish) and all fins are very heavily spotted with dark brown, and
the caudal fin is spotted with black. The markings and spots on the head are curved and almost vermiculate, with
the black mark over the eyeball and on the interorbital being curved, not always straight. The Tobago fish are
heavily blotched with brown so that the dorsal saddles are conspicuous, the black cheek bar is as nearly as broad as
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the pupil width and the marking above the pectoral fin base is very large and black. Fish from Belize are paler with
reduced spotting—about three rows of brown spots dorsally and the rest of the body whitish but with the lateral
vertical blotches on the body still visible. The dorsal fin spotting is less defined and the caudal fin is pale with only
a few brownish spots basally. Specimens from Atol das Rocas, Brazil, are very pale with no rows of dark spots on
the body or fins (which are unmarked); the fish resemble G. c a u e r e ns i s in colouring. Photos by Rob Myers of fish
from San Salvador (Fig. 26) show similar-coloured fish.
Ecology. Randall (1968) reported this species to be common, and found over “sand patches” at depths of
2–160 feet (0.5–48 m). We examined specimens collected from 1–36 m, from sand, coral, rock and rubble substrate
(usually sand or rubble).
Sponaugle and Cowen (1994) discussed the early larval life history and recruitment of G. t h o m p s o n i , which has
a long larval stage and is able to delay settlement. Gnatholepis thompsoni has a larval life of 59–122 (average 81.5)
days, and a post-settlement life of about 42 days, with a total lifespan of about 123 days (Shulman & Bermingham
1995; Thacker 2004a).
Kassi Cole made available to us her unpublished notes on the breeding biology of this species, made during her
research project on Coryphopterus at Puerto Rico:
“A nest is formed in a small depression in the sand, under the edge of a coral mound, or under a conch shell (or
in my case, under bricks that I set out for Coryphopterus glaucofraenum studies). The eggs are extremely small and
are attached to a thin, friable pad of sand which I suspect may be formed by the male with the use of secretions
from the accessory gonadal structure associated with the male reproductive system. Both the male and female are
associated with the nest, so there is biparental care. When both are at the nest site, often they are positioned with
the posterior half of their body under the cover of the overlying structure and the front half out in the open, and
often in a 'V' formation with respect to one another, so that their combined visual field is relatively wide.
Whenever I lifted up a brick, the guarding parent(s) would make vigorous side-to-side motions with the caudal
end of the body, then shoot away to about a distance of 2 m. At this point, nothing in the nest would be visible to
the naked eye. However, when I took a small dive knife and gently inserted it from the side, then moved the flat of
the blade back and forth while slowly lifting, the sand pad (or pieces of sand pad) would be lifted up, with the
attached eggs. So I concluded that the embryo-guarding parent(s) have a defense behavior that involves covering
them with sand before leaving the nest. Presumably, after the disturbance, they return and fan off the sand.
In every instance where I re-checked nests with eggs, the eggs, and guarding behavior, never lasted more than
two consecutive days. For example, when a brick was unoccupied one day, then occupied on the next day with
guarding fish and eggs, the eggs were present for two consecutive days, then disappeared. In several instances
where I removed the sand pad on a second day and placed it into a glass container to bring back to the lab, by the
time I got back to the lab the embryos had hatched, apparently due to mechanical stimulation. The newly hatched
larvae were extremely tiny. Therefore, I concluded that the embryonic developmental period was extremely short,
at around 48 h.” (Cole, in litt., May 2010).
Ross and Rhode (2004) reported two females collected in September from Onslow Bay, North Carolina, one of
which had eggs less than 0.1 mm diameter and a gonad index (gonad weight [body weight—gonad weight]
–1
) x
100) of 0.41; the other had mature eggs of 0.3 mm diameter and gonad index of 3.6.
Comparisons. This species looks very like G. c a u e re n s i s but differs in slightly in predorsal scale count (9–11,
modally 10, in G. t h o m p s o n i vs 7–12, modally 9, in G. c a u e re n s i s ), it has all the predorsal scales cycloid (vs always
cycloid in midline, with some ctenoid scales extending over the opercle in G. c a u e r e n s i s ), having the transverse
black line on the upper part of the eye joining the somewhat oblique to curved black line crossing cheek and
usually ending well behind end of jaw (vs transverse black bar on cheek mostly vertical to curved, ending past end
of jaw) and the dorsal half of the body with three to five rows of small dark spots (dorsal half of body usually with
2–3 rows of short lines in G. ca u e r en s i s ; some specimens with rows of spots).
Live specimens of G. c a u e r e n s i s from South Africa look almost identical in colour pattern to G. t h o m p s o n i , as
do heavily-spotted forms from dark substrates from Indonesia and Malaysia. The rows of spots along the side of
the body in G. t h o m p s o n i tend to be fairly even in intensity, while those in G. ca u e r e n s i s tend to have the dorsalmost
four or so rows darker than the remainder. These specimens would be practically impossible to identify from
photographs alone, without knowing their geographic origin.
Distribution. In the western Atlantic, known from southern Florida, North Carolina, the Bahamas and
Bermuda through to the Lesser Antilles (Sponaugle & Cole 1994); also reported from the oceanic islands of
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
Ascension, Madeira and St Helena, and Sao Tome Island, Cape Verde Islands and the Canary Islands in the eastern
Atlantic (Araujo & Freitas 2002).
Remarks. Rocha et al. (2005) presented a scenario, based on mtDNA, that Gnatholepis invaded the Atlantic
about 145,000 years ago from the Indian Ocean , and spread out to the central and eastern Atlantic 100,000 years
ago (and are continuing to invade the Canary Islands as sea temperatures rise). They considered G. t h o m p s o n i and
G. c a u e re n s i s (they referred to the latter as G. s c a p u l o s t i g m a ) to be sister taxa. Perhaps this is why Thacker (2004a:
579) refers to the Pacific and Caribbean populations of G. t h o m p s o n i being separated by the Isthmus of Panama.
We know of no Pacific populations of G. t h o m p s o n i .
TABLE 14. Measurements of specimens of Gnatholepis thompsoni, expressed as percentage of standard length (SL) or
head length (HL); n = 60.
Gnatholepis yoshinoi Suzuki and Randall, 2009
(Figs 27–29)
Gobius anjerensis—Kuiter and Tonozuka 2001: 671, figs A, B, D–F (Bali, Indonesia; Mabul, Malaysia).—Hayashi and
Shiratori 2003: 96, fig. 175 (Japan).
Gnatholepis yoshinoi Suzuki and Randall, 2009: 84–87 (Oura Bay, Okinawa, Ryukyu Islands, Japan).
Diagnosis. (From Suzuki and Randall 2009). A small Gnatholepis (up to 31 mm SL) with cycloid scales on head,
predorsal and pectoral fin base, distinct flap present at end of lower lip; first dorsal fin tall, pointed, with third spine
longest; distinctive black blotch on membrane behind first spine of first dorsal fin and one or more smaller black
spots on membrane between second and third spines; pectoral fin with rows of fine dark spots and speckles; second
dorsal and anal fin rays nearly always I,11; pectoral rays 15; lateral scales 27; about 7 cycloid predorsal scales.
Remarks. Specimens of this species were not available in time for examination (due to concatenation of
circumstances).
Means
(males)
Maximum
(males)
Minimum
(males)
Means
(females)
Maximum
(females)
Minimum
(females)
Standard length 37.4 58.0 25.0 35.5 46.0 21.5
Head length in SL 27.7 30.0 25.7 28.5 29.9 26.7
Head depth in HL 68.0 76.6 58.5 66.6 71.5 61.3
Head width in HL 68.4 77.2 57.3 66.5 71.6 59.4
Body depth at anus in SL 21.2 24.3 17.7 22.2 25.7 19.5
Body depth at D1 origin in SL 22.0 24.3 19.3 23.2 25.3 20.9
Caud. ped. length in SL 16.5 18.1 14.7 16.5 18.6 14.3
Caud. ped. depth in SL 11.6 13.8 10.3 11.6 13.2 10.2
Snout length in HL 31.1 38.2 25.7 30.6 35.2 25.7
Eye width in HL 26.8 32.0 23.1 27.3 35.5 23.0
Jaw length in HL 35.0 39.0 29.7 33.2 36.8 28.1
Interorbital in HL 8.0 11.0 4.9 7.8 9.8 6.0
Preorbital width in HL 19.1 23.0 13.6 19.8 24.8 16.7
Pectoral fin in SL 25.9 28.1 23.4 26.1 29.8 22.5
Pelvic fin in SL 27.5 31.0 24.7 26.7 30.0 23.3
Caudal fin in SL 32.4 37.4 29.8 30.6 32.2 26.6
Adpressed D1 in SL 25.5 31.4 21.2 25.0 29.5 20.9
3
rd
D1 fin spine in SL 18.0 21.6 15.8 17.9 20.9 16.1
4
th
D1 fin spine in SL 18.6 25.1 14.9 18.1 22.5 14.0
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FIGURE 27. Gnatholepis yoshinoi juvenile, Mabul, Malaysia. Photo by Rudie Kuiter.
FIGURE 28. Holotype of Gnatholepis yoshinoi, NSMT P.91420. Photograph by Toshiyuki Suzuki.
Figure 1 of the holotype and other photographs in Suzuki and Randall (2009: Figs 1–3) show the pelvic fins
having three bands of light and dark pigment (unlike the pattern on the pelvic fins of any other Gnatholepis
species), but the description does not mention this. This pattern of whitish and dark grey to blackish cross-bands on
the pelvic fins is shown on several photos of this species in the wild sent to us by Rudie Kuiter, and appears to be a
distinctive feature of this species (Figs 27–29).
Gnatholepis yoshinoi resembles G. a n j e re n s i s but the black blotches on the anterior part of the first dorsal fin
and row of large dark blotches along the lower side of the body and side of head are characteristic, as are the dark
and light cross-bands on the pelvic fins. The three known specimens show some differences in first dorsal fin
shape, with the fin varying from tall and rounded to very tall and pointed, as can be seen in published photographs
(Kuiter & Tonozuka 2001: 671, figs A, B, D–F; Suzuki & Randall 2009: figs 1–3) and in the holotype (Fig. 28). A
photograph of this species in Sydney Harbour, New South Wales, by Rudie Kuiter, shows a rounded fin and colour
pattern resembling that in Kuiter and Tonozuka (2001: 671, fig. E). We are not sure why Suzuki and Randall (2009)
did not refer to the other photographs of this species in Kuiter and Tonozuka (2001).
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A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
FIGURE 29. Probable Gnatholepis yoshinoi, Bali, Indonesia. Photo by Rudie Kuiter. Note banded pelvic fins, but no black
blotch on dorsal fin.
The fish in Figure 29 is unusual in that the first dorsal fin lacks the black blotches (and thus more resembles G.
anjerensis) but the pelvic fin can be seen as banded, which is not exhibited in G. a n j e re n s i s .
We consider that Hayashi and Shiratori’s (2003: 96) Figure 175 is of this species, although the tip of the first
dorsal fin is missing (looks like a piece has been taken out of it). However, the characteristic dark and light banding
on the pelvic fins is visible (through the transparent pectoral fin membrane), as is the smudgy dark cheek mark that
tends to cover the vertical black bar (and its other colour pattern features).
Incertae sedis
There is one taxon which may well be a Gnatholepis species, but it is uncertain if it is and to what species it could
belong. Eleotris feliceps Blyth, 1860, was described from Port Blair, Andaman Islands. The holotype might be at
ZSIC, but its exact whereabouts are not known. Day (1870) listed a specimen identified as this species from the
Museum in Calcutta, but he does not mention a type specimen. From the original description, it was possibly a
gobiid (eyes close together, mouth small, first dorsal fin spines slightly filamentous). Day (1870: 517) described
the head as being slightly adpressed, the anterior scales cycloid, posterior ctenoid, d2 I,10 A.11; scales 27; bands
extending ventrally from eye, with cheek and opercles scaled, suggesting Gnatholepis or Asterropteryx (or even
Callogobius).
The specimens in Zug et al. (1989) from Rotuma identified as Gnatholepis sp. were not borrowed due to time
constraints (four lots are present at USNM); therefore we cannot assign them to a species.
Acknowledgements
Our many thanks to Mark McGrouther (AMS), Mark Sabaj Pérez (ANSP), I-Shiung Chen (ASIZP), Arnold
Suzumoto and Jack Randall (BPBM); Dave Catania (CAS), Gavin Dally (NTM), Chris Thacker (LACM), Patrice
Pruvost (MNHN); Gento Shinohara (NSMT), Marcelo Kovacic (PMR), Vusi Mthombeni (SAIAB), Michael
Bougaardt (SAM) and Sandra Raredon (USNM) for loans of specimens. Dave Catania (CAS) kindly took data
from the holotype of G. t h o m p s o n i for us. Phil Willink (FMNH) sent us photographs of the holotype of G. c o r l e t t e i .
Toshiyuki Suzuki kindly gave us permission to use his photograph of the holotype of G. y o s h i n o i . Our thanks go to
Luis Rocha for information and discussion on Gnatholepis genetics, to Kassi Cole for information on breeding
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behaviour of Gnatholepis thompsoni and to Chris Thacker for her G. t h o m p s o n i suspensorium photo.
Gerry Allen, Ole Brett, Sergey Bogorodsky, Phil Heemstra, Doug Hoese, Dennis King, Rudie Kuiter, Jeff Leis,
Rob Myers, Robert Patzner, Jack Randall, Sandra Raredon, Dick Wass and Rick Winterbottom kindly made
available to us many images of living and freshly dead Gnatholepis species, to help us work out which species
were what—their willingness to help in this way was invaluable.
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Zootaxa 3529 © 2012 Magnolia Press · 63
A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
APPENDIX 1. List of all Gnatholepis specimens, and the data, used in the canonical analysis of Principal Co–ordinates
Analysis (CAP).
Sample ID
(catalogue number
Species Sex SL (mm) Location
Counts and proportions used in CAP analysis
Pectoral right
Pectoral left
Lat.scales
TRB
Pred.scales
HL in SL
HD in HL
HW in HL
BDA in SL
BD D1 in SL
CPL in SL
CPD in SL
Snout in HL
Eye in HL
Jaw in HL
Preorb. in HL
AMS I.21874–001 G. a n j e r e n s i s male 38.0 Moorea, Society Islands 16.0 16.0 27.0 10.0 10.0 26.6 72.2 69.8 23.0 25.8 16.1 13.2 31.3 26.8 39.3 22.3
AMS I.21874–001 G. a n j e r e n s i s female 35.0 Moorea, Society Islands 17.0 17.0 27.0 10.5 10.0 30.0 70.5 69.5 24.9 14.6 17.1 12.3 31.4 29.5 35.2 21.9
AMS I.28999–010 G. a n j e r e n s i s male 31.5 Cocos–Keeling Islands 16.0 15.0 27.0 10.0 9.0 29.5 67.7 66.7 20.6 22.2 17.5 11.1 35.5 28.0 34.4 21.5
AMS I.28999–010 G. a n j e r e n s i s female 36.5 Cocos–Keeling Islands 16.0 16.0 28.0 11.5 10.0 29.6 68.5 65.7 21.1 23.0 18.1 10.7 33.3 26.9 33.3 24.1
AMS I.30940–006 G. a n j e r e n s i s male 37.0 Moorea, Society Islands 15.0 16.0 26.0 11.5 10.0 29.5 69.7 67.9 22.7 23.2 15.7 11.4 36.7 25.7 34.9 21.1
AMS I.30940–006 G. a n j e r e n s i s male 34.0 Moorea, Society Islands 16.0 16.0 27.0 10.5 10.0 30.3 69.9 73.8 22.9 23.8 12.9 11.8 40.8 29.1 35.0 21.4
AMS I.30940–006 G. a n j e r e n s i s male 31.0 Moorea, Society Islands 16.0 16.0 26.0 10.5 10.0 29.7 67.4 67.4 21.6 24.5 15.5 11.3 34.8 29.3 40.2 18.5
AMS I.30940–006 G. a n j e r e n s i s male 30.0 Moorea, Society Islands 16.0 15.0 25.0 10.0 10.0 30.3 70.3 71.4 22.7 25.3 15.7 12.0 34.1 29.7 39.6 19.8
AMS I.30940–006 G. a n j e r e n s i s female 33.0 Moorea, Society Islands 16.0 16.0 26.0 10.5 10.0 30.6 71.3 69.3 23.3 24.8 16.1 11.5 36.6 29.7 34.7 21.8
AMS I.30940–006 G. a n j e r e n s i s female 26.5 Moorea, Society Islands 16.0 16.0 27.0 10.0 10.0 31.7 67.9 70.2 23.4 24.5 18.5 12.1 35.7 27.4 32.1 21.4
AMS I.35852–003 G. a n j e r e n s i s female 65.0 One Tree Island 16.0 16.0 26.0 10.5 10.0 27.8 71.8 70.2 23.8 24.2 16.6 12.5 36.5 26.0 32.0 23.8
AMS I.40826–001 G. a n j e r e n s i s male 44.0 Guam, Mariana Islands 16.0 16.0 27.0 10.5 9.0 29.5 71.5 64.6 23.4 23.6 15.0 12.0 36.9 26.2 36.9 22.3
BPBM 26651 G. a n j e r e n s i s female 31.0 Neotype Gobius
anjerensis
16.0 16.0 26.0 9.0 7.0 30.0 66.7 67.7 22.3 22.6 14.8 11.3 34.4 30.1 34.4 21.5
BPBM 267 G. a n j e r e n s i s male 44.5 Syntype Gobius
deltoides
16.0 16.0 27.0 10.5 11.0 29.2 75.4 57.7 22.5 23.1 14.4 11.5 33.8 25.4 38.5 20.8
BPBM 267 G. a n j e r e n s i s male 37.5 Syntype Gobius
deltoides
15.0 16.0 28.0 10.5 10.0 28.5 71.0 60.7 22.4 22.7 14.7 12.0 32.7 28.0 37.4 23.4
BPBM 34045 G. a n j e r e n s i s male 45.0 Johnston Atoll 17.0 17.0 27.0 10.0 9.0 26.7 75.0 70.8 21.3 21.8 16.0 11.6 34.2 25.0 36.7 20.8
BPBM 34045 G. a n j e r e n s i s male 32.5 Johnston Atoll 18.0 18.0 26.0 9.0 9.0 28.9 63.8 66.0 21.5 22.8 16.3 11.4 31.9 24.5 35.1 22.3
BPBM 34045 G. a n j e r e n s i s female 27.0 Johnston Atoll 18.0 18.0 27.0 9.0 9.0 28.5 68.8 66.2 19.6 21.1 16.7 10.4 35.1 27.3 33.8 22.1
CAS 51530 G. a n j e r e n s i s male 43.0 Taravao, Tahiti 15.0 15.0 27.0 11.0 8.0 29.3 75.4 68.3 23.3 22.6 17.4 11.2 32.5 23.8 35.7 23.8
CAS 51530 G. a n j e r e n s i s female 49.0 Taravao, Tahiti 16.0 16.0 28.0 11.0 9.0 30.6 70.7 62.7 24.5 25.1 16.1 11.2 33.3 24.0 36.7 23.3
CAS 51530 G. a n j e r e n s i s female 41.5 Taravao, Tahiti 15.0 15.0 26.0 10.0 8.0 28.4 73.7 64.4 23.1 24.8 15.4 10.8 31.4 30.5 38.1 22.0
ex USNM 308212 G. a n j e r e n s i s female 56.0 One Tree Island 17.0 17.0 27.0 11.0 8.0 28.9 71.6 72.2 23.8 24.6 18.6 12.1 37.0 25.9 34.6 26.5
NSMT P.64544 G. a n j e r e n s i s male 27.5 Ambon 15.0 15.0 26.0 10.0 9.0 29.8 72.0 67.1 22.2 23.6 18.2 11.6 30.5 28.0 35.4 20.7
NSMT P.64544 G. a n j e r e n s i s female 27.0 Ambon 16.0 16.0 25.0 10.0 8.0 33.3 65.6 63.3 23.7 24.8 16.3 11.9 33.3 27.8 35.6 18.9
NSMT P.64544 G. a n j e r e n s i s female 26.0 Ambon 16.0 15.0 27.0 10.0 9.0 30.4 62.0 64.6 21.9 22.7 16.5 11.2 32.9 27.8 32.9 20.3
NTM S.11973–037 G. a n j e r e n s i s male 32.0 Ashmore Reef, Indian
Ocean
15.0 15.0 26.0 10.5 9.0 30.0 72.9 72.9 22.5 15.0 15.9 11.9 37.5 27.1 34.4 21.9
NTM S.11973–037 G. a n j e r e n s i s female 37.5 Ashmore Reef, Indian
Ocean
16.0 16.0 27.0 11.0 9.0 31.2 82.1 78.6 24.8 15.5 16.0 12.3 44.4 26.5 33.3 23.1
NTM S.11983–002 G. a n j e r e n s i s male 33.5 Ashmore Reef, Indian
Ocean
16.0 17.0 26.0 10.0 9.0 30.4 74.5 74.5 23.0 14.3 16.1 11.9 39.2 26.5 33.3 23.5
NTM S.13965–005 G. a n j e r e n s i s female 45.5 Kenya 17.0 17.0 26.0 11.0 9.0 31.0 78.0 75.2 26.6 15.8 15.4 13.0 37.6 24.8 34.0 24.8
SAIAB 19636 G. a n j e r e n s i s male 47.0 Mozambique 15.0 16.0 28.0 11.5 9.0 28.1 69.7 68.9 22.1 23.4 15.5 10.9 35.6 29.5 34.8 22.7
SAIAB 19636 G. a n j e r e n s i s female 52.0 Mozambique 15.0 15.0 26.0 9.0 8.0 29.8 76.8 67.7 24.0 27.1 15.4 11.7 35.5 28.4 36.1 22.6
SAIAB 19636 G. a n j e r e n s i s female 41.0 Mozambique 16.0 16.0 28.0 10.0 10.0 28.8 67.8 62.7 22.4 22.4 15.6 11.0 33.1 28.0 36.4 23.7
SAIAB 2112 G. a n j e r e n s i s male 51.0 Mauritius, Indian Ocean 16.0 16.0 27.0 11.5 11.0 29.4 80.0 78.7 23.9 25.5 16.7 12.4 42.0 24.7 40.0 22.0
SAIAB 2112 G. a n j e r e n s i s male 46.5 Mauritius, Indian Ocean 16.0 15.0 27.0 11.0 9.0 27.7 83.7 82.2 23.2 23.9 17.2 11.8 43.4 29.5 41.1 22.5
SAIAB 2112 G. a n j e r e n s i s female 51.0 Mauritius 16.0 16.0 28.0 11.0 10.0 29.8 74.3 74.3 22.9 24.7 16.3 11.4 40.1 26.3 36.2 21.7
SAIAB 2317 G. a n j e r e n s i s female 45.5 Mauritius 17.0 16.0 27.0 11.5 10.0 29.0 81.8 79.5 24.8 27.7 18.2 11.9 37.1 30.3 34.8 25.0
SAIAB 2317 G. a n j e r e n s i s female 42.5 Mauritius 14.0 15.0 26.0 10.0 9.0 27.5 75.2 72.6 24.7 25.9 16.9 11.8 34.2 28.2 34.2 21.4
SAIAB 2576 G. a n j e r e n s i s male 84.0 Thompsons Bay, KZN,
South Africa
16.0 16.0 27.0 12.0 13.0 28.2 74.7 65.8 26.3 25.2 14.5 13.8 40.5 23.2 38.0 27.8
SAIAB 70332 G. a n j e r e n s i s male 51.5 Rodrigues 17.0 17.0 27.0 10.0 8.0 27.2 75.0 64.3 23.5 24.1 16.7 13.0 32.1 25.7 37.1 18.6
USNM 109399 G. a n j e r e n s i s male 39.0 Paratype Gobius
deltoides
16.0 16.0 27.0 10.5 10.0 31.0 66.9 53.7 23.6 23.6 15.4 12.3 33.1 29.8 36.4 24.0
USNM 211025 G. a n j e r e n s i s male 34.0 Moluccas 16.0 15.0 26.0 11.0 9.0 29.1 69.7 68.7 23.8 23.8 15.0 11.8 30.3 26.3 36.4 18.2
USNM 211025 G. a n j e r e n s i s male 32.5 Moluccas 15.0 15.0 25.0 10.0 9.0 30.2 66.3 67.3 22.8 24.0 15.1 11.7 22.4 26.5 34.7 21.4
...... continued on the next page
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64 · Zootaxa 3529 © 2012 Magnolia Press
APPENDIX 1. (Continued)
Sample ID
(catalogue number
Species Sex SL (mm) Location
Counts and proportions used in CAP analysis
Pectoral right
Pectoral left
Lat.scales
TRB
Pred.scales
HL in SL
HD in HL
HW in HL
BDA in SL
BD D1 in SL
CPL in SL
CPD in SL
Snout in HL
Eye in HL
Jaw in HL
Preorb. in HL
USNM 211025 G. a n j e r e n s i s male 34.0 Moluccas 16.0 16.0 26.0 10.0 9.0 30.6 65.4 66.3 24.4 24.7 16.8 11.8 31.7 23.1 34.6 22.1
USNM 211025 G. a n j e r e n s i s female 30.5 Moluccas 16.0 16.0 26.0 10.0 8.0 31.5 65.6 64.6 24.6 26.6 15.7 12.1 30.2 26.0 34.4 20.8
USNM 211025 G. a n j e r e n s i s female 32.0 Moluccas 16.0 16.0 26.0 10.0 9.0 30.0 63.5 67.7 22.8 23.1 14.7 11.3 29.2 26.0 34.4 20.8
USNM 313779 G. a n j e r e n s i s male 43.0 Egypt, Red Sea 16.0 16.0 26.0 11.5 8.0 28.8 67.7 66.9 23.0 24.2 16.0 11.4 33.1 27.4 35.5 22.6
USNM 327464 G. a n j e r e n s i s male 43.5 Zubair I., Red Sea 16.0 17.0 27.0 10.5 10.0 29.4 70.3 68.0 23.7 24.8 14.7 12.0 35.9 27.3 35.9 25.8
USNM 327464 G. a n j e r e n s i s male 48.5 Zubair I., Red Sea 17.0 16.0 27.0 11.0 9.0 28.2 70.8 67.2 23.1 23.7 15.9 11.8 36.5 27.0 38.0 24.1
USNM 327464 G. a n j e r e n s i s male 45.5 Zubair I., Red Sea 16.0 16.0 27.0 10.5 9.0 28.6 74.6 67.7 24.2 25.5 15.4 12.7 35.4 27.7 41.5 26.2
USNM 327464 G. a n j e r e n s i s male 44.0 Zubair I., Red Sea 17.0 16.0 27.0 10.5 8.0 29.1 68.8 72.7 23.0 24.5 15.0 12.5 35.9 27.3 39.8 21.9
USNM 327464 G. a n j e r e n s i s female 29.5 Zubair I., Red Sea 15.0 16.0 26.0 10.5 8.0 29.5 69.0 62.1 23.7 25.1 18.0 11.9 31.0 29.9 33.3 18.4
USNM 327464 G. a n j e r e n s i s female 31.0 Zubair I., Red Sea 15.0 16.0 26.0 10.0 8.0 29.0 66.7 65.6 23.2 23.9 16.1 11.3 31.1 30.0 35.6 21.1
BPBM 13401 G. ca u d im a c ul a t a female 34.5 Gulf of Aqaba 15.0 15.0 27.0 8.0 6.0 29.0 70.0 65.0 22.3 23.8 16.8 10.7 35.0 26.0 37.0 19.0
BPBM 18231 G. ca u d im a c ul a t a male 33.0 Gulf of Aqaba 16.0 16.0 28.0 10.0 9.0 26.6 69.9 64.6 23.1 23.0 17.0 10.6 32.2 30.0 36.7 20.0
BPBM 18231 G. ca u d im a c ul a t a male 32.5 Gulf of Aqaba 15.0 15.0 26.0 11.0 8.0 27.3 73.3 68.9 21.8 22.5 17.5 11.1 37.4 30.8 34.1 19.8
BPBM 18231 G. ca u d im a c ul a t a male 35.0 Gulf of Aqaba 16.0 16.0 26.0 10.0 9.0 27.7 73.2 69.1 21.7 24.9 16.6 11.4 33.0 28.9 35.1 20.6
BPBM 18231 G. ca u d im a c ul a t a male 37.0 Gulf of Aqaba 16.0 16.0 26.0 10.0 8.0 27.6 82.4 74.5 22.2 23.5 16.8 11.6 38.2 30.4 35.3 21.6
BPBM 18231 G. ca u d im a c ul a t a female 32.0 Gulf of Aqaba 16.0 16.0 27.0 10.0 8.0 28.8 67.4 65.2 22.5 25.6 19.1 11.3 32.6 32.6 37.0 21.7
BPBM 18231 G. ca u d im a c ul a t a female 26.0 Gulf of Aqaba 16.0 16.0 27.0 9.0 9.0 30.0 70.5 66.7 23.1 25.0 17.3 10.4 30.8 30.8 32.1 20.5
BPBM 27442 G. ca u d im a c ul a t a male 28.0 Sudan 15.0 17.0 26.0 9.5 8.0 30.7 62.8 55.8 23.9 23.9 16.8 11.1 27.9 29.1 33.7 15.1
BPBM 30457 G. ca u d im a c ul a t a male 33.3 Persian Gulf 16.0 16.0 26.0 11.0 8.0 30.0 68.0 68.0 20.1 22.5 19.5 11.1 37.0 27.0 34.0 21.0
BPBM 30457 G. ca u d im a c ul a t a male 35.0 Persian Gulf 16.0 16.0 28.0 11.0 9.0 29.1 65.7 59.8 20.3 21.4 17.4 10.9 31.4 26.5 35.3 19.6
BPBM 30457 G. ca u d im a c ul a t a male 34.0 Persian Gulf 16.0 16.0 28.0 10.0 9.0 28.8 64.3 60.2 20.9 21.2 16.5 10.6 30.6 26.5 36.7 20.4
BPBM 33350 G. ca u d im a c ul a t a male 36.5 Saudi Arabia 15.0 15.0 27.0 10.0 7.0 27.7 61.4 58.4 19.2 19.5 17.8 10.4 28.7 25.7 35.6 15.8
BPBM 33350 G. ca u d im a c ul a t a male 30.0 Saudi Arabia 16.0 16.0 27.0 9.0 8.0 29.7 62.9 59.6 21.3 22.7 17.7 11.0 32.6 25.8 34.8 19.1
BPBM 33373 G. ca u d im a c ul a t a male 42.5 Saudi Arabia 16.0 16.0 26.0 10.5 8.0 26.6 69.9 64.6 23.1 23.3 16.7 11.8 34.5 26.5 39.8 18.6
BPBM 33374 G. ca u d im a c ul a t a female 36.5 Saudi Arabia 17.0 17.0 26.0 10.0 8.0 29.6 66.7 59.3 24.4 23.0 16.4 11.5 31.5 25.0 33.3 19.4
ex–USNM 327464 G. c a ud i m ac u l a ta male 27.0 Zubair Island, Persian
Gulf
16.0 16.0 26.0 10.0 8.0 27.4 73.0 70.3 21.9 23.3 17.4 11.9 29.7 29.7 36.5 17.6
ex–USNM 327464 G. c a ud i m ac u l a ta female 28.0 Zubair Island, Persian
Gulf
15.0 15.0 26.0 10.0 7.0 29.6 69.9 65.1 24.6 24.6 16.4 11.4 27.7 30.1 36.1 15.7
PMR VP.2219 G. ca u d i m ac u l a ta female 30.5 Sudan 16.0 16.0 26.0 10.0 7.0 24.6 78.7 76.0 19.0 21.0 14.4 10.5 33.3 29.3 37.3 20.0
PMR VP.2220 G. ca u d i m ac u l a ta male 28.5 Sudan 16.0 16.0 25.0 10.0 9.0 28.4 70.4 70.4 21.1 22.8 16.8 10.9 34.6 28.4 33.3 18.5
USNM 313780 G. c au d im a c ul a t a male 29.0 Ethiopia Red Sea 15.0 15.0 26.0 10.0 8.0 27.2 73.4 68.4 13.4 22.1 17.9 11.4 30.4 29.1 35.4 16.5
USNM 313780 G. c au d im a c ul a t a male 28.5 Ethiopia Red Sea 16.0 16.0 26.0 10.0 8.0 29.1 65.1 65.1 22.5 21.8 15.1 11.2 31.3 27.7 34.9 18.1
USNM 313780 G. c au d im a c ul a t a female 28.5 Ethiopia Red Sea 16.0 15.0 26.0 10.0 9.0 29.8 71.8 72.9 22.5 24.2 15.8 11.2 32.9 25.9 36.5 16.5
USNM 313780 G. c au d im a c ul a t a female 29.0 Ethiopia Red Sea 16.0 16.0 26.0 11.0 8.0 28.6 71.1 71.1 23.8 24.1 17.9 11.4 28.9 30.1 37.3 18.1
USNM 313780 G. c au d im a c ul a t a female 28.5 Ethiopia Red Sea 15.0 16.0 26.0 10.0 8.0 28.4 75.3 72.8 23.5 24.2 16.5 11.6 30.9 30.9 39.5 18.5
USNM 313780 G. c au d im a c ul a t a female 28.5 Ethiopia Red Sea 16.0 16.0 26.0 10.0 8.0 28.8 69.5 63.4 23.5 23.9 16.8 11.2 29.3 29.3 32.9 14.6
USNM 327425 G. c au d im a c ul a t a male 41.0 HOLOTYPE 15.0 15.0 26.0 10.0 8.0 26.8 68.2 70.0 22.2 22.2 17.6 10.7 35.5 27.3 38.2 20.9
USNM 327425 G. c au d im a c ul a t a male 36.0 Israel, Red Sea 16.0 16.0 27.0 10.0 8.0 26.4 74.7 70.5 24.2 24.2 15.6 11.7 32.6 29.5 33.7 18.9
USNM 327425 G. c au d im a c ul a t a male 33.0 Israel, Red Sea 16.0 16.0 25.0 9.5 8.0 28.2 68.8 73.1 23.9 25.5 17.0 10.9 33.3 28.0 36.6 21.5
USNM 327425 G. c au d im a c ul a t a female 33.5 Israel, Red Sea 16.0 16.0 26.0 9.5 9.0 28.4 69.5 69.5 23.0 25.7 16.4 10.7 31.6 28.4 37.9 21.1
AMS I.17445–049 G. c a u e r e n s i s female 42.5 One Tree I, Qld 16.0 16.0 27.0 10.0 10.0 26.4 67.0 69.6 21.9 22.4 14.6 12.0 29.5 26.8 35.7 20.5
AMS I.19500–003 G. c a u e r e n s i s female 31.5 Parsley Bay, NSW 16.0 16.0 26.0 10.0 10.0 27.9 68.2 69.3 13.7 21.3 16.5 12.4 36.4 27.3 34.1 22.7
AMS I.19641–022 G. c a u e r e n s i s male 37.5 Tantabiddi Ck, WA 16.0 16.0 27.0 9.5 8.0 29.6 68.5 72.1 21.3 22.7 16.5 11.7 36.9 25.2 35.1 23.4
AMS I.19641–022 G. c a u e r e n s i s male 40.0 Tantabiddi Ck, WA 17.0 16.0 26.0 10.0 9.0 27.8 73.9 77.5 21.3 21.8 15.3 12.0 38.7 26.1 37.8 23.4
AMS I.20757–072 G. c a u e r e n s i s male 36.0 Raine Island, Great
Barrier Reef
17.0 17.0 26.0 10.0 9.0 26.9 64.9 64.9 20.6 20.3 17.5 11.1 28.9 27.8 32.0 16.5
AMS I.20757–072 G. c a u e r e n s i s male 28.0 Raine Island, N Great
Barrier Reef
17.0 17.0 26.0 9.0 9.0 28.9 66.7 65.4 21.4 23.6 17.5 11.4 25.9 29.6 30.9 18.5
...... continued on the next page
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Zootaxa 3529 © 2012 Magnolia Press · 65
A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
APPENDIX 1. (Continued)
Sample ID
(catalogue number
Species Sex SL (mm) Location
Counts and proportions used in CAP analysis
Pectoral right
Pectoral left
Lat.scales
TRB
Pred.scales
HL in SL
HD in HL
HW in HL
BDA in SL
BD D1 in SL
CPL in SL
CPD in SL
Snout in HL
Eye in HL
Jaw in HL
Preorb. in HL
AMS I.20757–072 G. c a u e r e n s i s female 27.5 Raine Island, Great
Barrier Reef
16.0 16.0 26.0 10.0 9.0 29.8 63.4 64.6 23.6 24.7 16.4 11.6 28.0 28.0 30.5 17.1
AMS I.20757–072 G. c a u e r e n s i s female 30.5 Raine Island, Great
Barrier Reef
16.0 16.0 27.0 10.0 8.0 27.9 76.5 69.4 23.3 24.6 18.0 11.5 25.9 27.1 27.1 17.6
AMS I.20774–107 G. c a u e r e n s i s male 39.0 Cape Melville,
Queensland
18.0 17.0 26.0 10.0 8.0 27.4 74.8 78.5 23.8 23.8 17.2 12.3 38.3 28.0 36.4 20.6
AMS I.20774–107 G. c a u e r e n s i s male 40.0 Cape Melville,
Queensland
16.0 16.0 26.0 10.0 9.0 27.0 70.4 69.4 21.8 23.0 16.5 11.8 32.4 25.9 38.0 19.4
AMS I.20774–107 G. c a u e r e n s i s male 28.0 Cape Melville,
Queensland
16.0 16.0 26.0 10.0 7.0 27.9 67.9 65.4 21.4 22.5 16.4 10.7 32.1 26.9 32.1 20.5
AMS I.20774–107 G. c a u e r e n s i s female 34.0 Cape Melville,
Queensland
16.0 16.0 27.0 10.0 8.0 27.9 67.4 68.4 22.1 23.2 15.6 11.5 36.8 27.4 35.8 31.6
AMS I.21907–003 G. c a u e r e n s i s male 34.0 Philippines 18.0 17.0 27.0 11.5 9.0 28.8 66.3 68.4 22.9 23.2 16.8 12.4 30.6 27.6 36.7 19.4
AMS I.21907–003 G. c a u e r e n s i s female 31.5 Philippines 17.0 12.0 27.0 10.0 9.0 28.9 67.0 67.0 22.5 24.1 16.5 11.4 27.5 27.5 35.2 17.6
AMS I.23492–015 G. c a u e r e n s i s male 34.5 Miyakejima, S Japan 17.0 17.0 28.0 9.0 9.0 27.5 71.6 70.5 22.6 23.5 15.1 12.2 32.6 27.4 31.6 21.1
AMS I.27138–055 G. c a u e r e n s i s male 44.0 Middleton Reef, off E
Australia
17.0 17.0 27.0 10.0 9.0 26.1 68.7 65.2 21.4 22.3 15.5 11.6 33.0 24.3 27.8 21.7
AMS I.27138–055 G. c a u e r e n s i s female 35.5 Middleton Reef, NSW 16.0 15.0 27.0 10.0 8.0 27.6 66.3 69.4 21.7 22.3 14.6 11.3 32.7 26.5 34.7 20.4
AMS I.27138–055 G. c a u e r e n s i s female 30.0 Middleton Reef, NSW 17.0 16.0 26.0 10.0 10.0 28.7 69.8 72.1 22.7 24.3 16.7 11.3 34.9 27.9 37.2 18.6
AMS I.27142–030 G. c a u e r e n s i s male 49.0 Middleton Reef, off E
Australia
15.0 15.0 26.0 10.0 8.0 26.9 66.7 67.4 23.1 21.0 13.1 12.4 30.3 23.5 34.1 20.5
AMS I.27142–030 G. c a u e r e n s i s male 56.0 Middleton Reef, off E
Australia
16.0 17.0 27.0 10.0 9.0 27.5 73.4 71.4 21.6 22.0 13.9 12.0 39.0 20.1 34.4 21.4
AMS I.27142–030 G. c a u e r e n s i s male 33.0 Middleton Reef, off E
Australia
16.0 16.0 28.0 10.0 7.0 28.8 64.2 63.2 21.8 22.1 18.2 12.1 33.7 27.4 33.7 22.1
AMS I.27142–030 G. c a u e r e n s i s female 51.0 Middleton Reef, off E
Australia
16.0 15.0 29.0 10.5 9.0 27.8 71.1 66.9 22.4 23.5 11.4 17.5 35.2 26.8 35.2 23.2
AMS I.27142–030 G. c a u e r e n s i s female 40.5 Middleton Reef, off E
Australia
16.0 16.0 25.0 10.0 8.0 26.9 67.9 64.2 24.7 24.7 15.6 13.3 33.0 25.7 34.9 22.9
AMS I.27156–035 G. c a u e r e n s i s male 53.0 Elizabeth Reef, off E
Australia
16.0 16.0 24.0 10.0 11.0 26.6 70.9 73.0 22.8 22.3 15.1 12.1 35.5 25.5 37.6 21.3
AMS I.27156–035 G. c a u e r e n s i s male 49.0 Elizabeth Reef, off E
Australia
16.0 15.0 26.0 10.0 10.0 27.3 67.2 63.4 22.2 23.5 15.5 12.4 36.6 23.9 35.1 20.9
AMS I.27156–035 G. c a u e r e n s i s female 42.5 Elizabeth Reef, off E
Australia
16.0 15.0 27.0 10.0 11.0 28.9 65.0 61.8 22.4 24.7 15.8 11.3 35.0 26.8 35.0 20.3
AMS I.33749–122 G. c a u e r e n s i s male 33.0 Boot Reef, Coral Sea 16.0 15.0 26.0 10.0 9.0 27.3 64.4 63.3 20.6 22.1 15.8 11.2 31.1 27.8 32.2 20.0
AMS I.33749–122 G. c a u e r e n s i s female 29.5 Boot Reef, Coral Sea 16.0 16.0 26.0 10.0 9.0 27.8 64.6 65.9 21.7 22.4 17.6 11.5 26.8 28.0 35.4 19.5
AMS I.40155–059 G. c a u e r e n s i s male 36.0 Ambulong, PhiIippines 17.0 16.0 27.0 10.5 8.0 26.9 74.2 75.3 21.1 19.4 15.8 11.1 35.1 27.8 36.1 19.6
AMS IB.4004 G. c a u e r e n s i s female 42.5 Heron Island, Great
Barrier Reef
16.0 16.0 27.0 10.0 9.0 28.0 63.0 68.1 21.2 22.8 15.8 11.8 28.6 26.9 35.3 21.0
BPBM 11003 G. c a u e r e n s i s male 42.0 Marquesas Islands, S
Pacific
18.0 18.0 27.0 10.0 9.0 27.9 65.0 66.7 19.8 19.8 16.2 12.1 35.0 26.5 36.8 19.7
BPBM 11869 G. c a u e r e n s i s male 53.0 Marquesas Islands, S
Pacific
18.0 18.0 28.0 10.5 9.0 25.7 66.2 68.4 19.8 19.2 17.5 11.3 33.1 27.2 37.5 19.1
BPBM 11869 G. c a u e r e n s i s male 49.5 Marquesas Islands, S
Pacific
17.0 17.0 27.0 10.5 9.0 26.3 65.4 66.9 20.4 21.8 15.2 12.1 35.4 26.2 36.9 20.0
BPBM 11869 G. c a u e r e n s i s male 37.0 Marquesas Islands, S
Pacific
17.0 17.0 29.0 11.0 8.0 26.2 62.9 64.9 18.4 18.6 14.9 10.8 32.0 26.8 38.1 19.6
BPBM 11869 G. c a u e r e n s i s male 32.5 Marquesas Islands, S
Pacific
17.0 17.0 27.0 10.0 8.0 28.0 68.1 73.6 20.6 21.5 16.9 11.1 35.2 27.5 35.2 18.7
BPBM 11869 G. c a u e r e n s i s female 41.5 Marquesas Islands, S
Pacific
18.0 18.0 27.0 10.5 8.0 28.4 65.3 74.6 21.2 21.7 16.4 11.3 38.1 24.6 36.4 20.3
BPBM 12621 G. c a u e r e n s i s female 38.0 Marquesas Islands, S
Pacific
18.0 18.0 27.0 12.0 9.0 28.4 63.9 64.8 22.1 22.4 15.0 11.8 32.4 25.0 35.2 18.5
BPBM 15131 G. c a u e r e n s i s male 36.0 Maui 17.0 16.0 28.0 10.0 9.0 25.8 69.9 66.7 22.5 21.7 14.7 11.9 29.0 26.9 37.6 17.2
BPBM 15131 G. c a u e r e n s i s male 24.5 Maui 17.0 17.0 27.0 10.0 10.0 26.1 68.8 67.2 20.0 21.2 16.7 10.6 29.7 28.1 35.9 18.8
...... continued on the next page
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LARSON & BUCKLE
66 · Zootaxa 3529 © 2012 Magnolia Press
APPENDIX 1. (Continued)
Sample ID
(catalogue number
Species Sex SL (mm) Location
Counts and proportions used in CAP analysis
Pectoral right
Pectoral left
Lat.scales
TRB
Pred.scales
HL in SL
HD in HL
HW in HL
BDA in SL
BD D1 in SL
CPL in SL
CPD in SL
Snout in HL
Eye in HL
Jaw in HL
Preorb. in HL
BPBM 15131 G. c a u e r e n s i s male 37.0 Maui 17.0 17.0 28.0 9.5 9.0 25.7 70.5 68.4 22.4 20.8 16.8 10.8 29.5 27.4 36.8 17.9
BPBM 15131 G. c a u e r e n s i s male 39.0 Maui 17.0 16.0 27.0 10.5 9.0 24.6 71.9 69.8 22.3 20.0 15.9 11.3 32.3 28.1 34.4 19.8
BPBM 15131 G. c a u e r e n s i s male 39.0 Maui 17.0 17.0 28.0 9.5 10.0 24.1 78.7 69.1 23.6 21.8 15.6 11.5 33.0 27.7 35.1 20.2
BPBM 15131 G. c a u e r e n s i s female 31.5 Maui 16.0 16.0 27.0 10.0 9.0 27.3 69.8 68.6 23.2 23.2 17.5 10.8 30.2 26.7 33.7 17.4
BPBM 15131 G. c a u e r e n s i s female 35.0 Maui 17.0 17.0 27.0 10.5 8.0 27.4 67.7 65.6 22.0 22.3 18.0 11.1 29.2 26.0 33.3 17.7
BPBM 15131 G. c a u e r e n s i s female 41.0 Maui 17.0 17.0 28.0 9.0 9.0 26.8 66.4 72.7 22.0 20.5 14.6 10.5 37.3 23.6 37.3 22.7
BPBM 15131 G. c a u e r e n s i s female 33.5 Maui 17.0 17.0 27.0 10.0 10.0 27.2 71.4 71.4 21.5 22.7 15.5 10.4 33.0 27.5 34.1 19.8
BPBM 15131 G. c a u e r e n s i s female 30.5 Maui 17.0 17.0 27.0 9.5 10.0 27.2 68.7 63.9 23.0 24.3 16.7 10.2 30.1 26.5 31.3 20.5
BPBM 16846 G. c a u e r e n s i s male 41.5 Pitcairn Island 18.0 16.0 26.0 10.0 9.0 26.3 73.4 78.0 20.5 20.5 14.7 12.0 37.6 24.8 39.4 21.1
BPBM 17267 G. c a u e r e n s i s female 38.0 Rapa 18.0 18.0 27.0 10.5 10.0 27.9 69.8 70.8 21.3 22.1 17.6 11.1 37.7 26.4 33.0 20.8
BPBM 17276 G. c a u e r e n s i s female 41.0 Holotype G. c a u e r e n s i s
australis
19.0 19.0 27.0 10.0 9.0 26.8 70.9 66.4 20.7 22.2 17.1 12.2 31.8 24.5 33.6 19.1
BPBM 37861 G. c a u e r e n s i s female 38.5 Kona coast 17.0 17.0 28.0 10.0 9.0 27.5 70.8 67.9 24.7 24.9 17.9 12.2 30.2 29.2 35.8 20.8
BPBM 37861 G. c a u e r e n s i s female 29.5 Kona coast 17.0 17.0 29.0 10.5 9.0 25.8 67.1 65.8 21.7 21.7 16.6 11.2 28.9 26.3 35.5 19.7
BPBM 38376 G. c a u e r e n s i s male 35.5 Rapa 18.0 18.0 27.0 9.0 8.0 27.3 69.1 73.2 21.4 22.5 15.5 11.5 37.1 25.8 34.0 22.7
MNHN 1980–744 G. c a u e r e n s i s female 36.0 New Caledonia 17.0 16.0 26.0 10.5 9.0 26.9 68.0 59.8 22.8 20.6 16.7 11.9 29.9 27.8 30.9 16.5
MNHN 1980–94 G. c a u e r e n s i s male 32.5 Gambier Island, S
Pacific
17.0 16.0 28.0 10.5 11.0 28.0 65.9 60.4 20.9 22.8 15.4 11.1 34.1 27.5 35.2 19.8
NSMT P.70196 G. c a u e r e n s i s male 30.0 Lombok 17.0 16.0 27.0 10.0 8.0 30.7 64.1 60.9 18.7 22.0 16.3 12.0 31.5 29.3 34.8 16.3
NSMT P.70196 G. c a u e r e n s i s female 35.0 Lombok 13.0 17.0 27.0 10.5 8.0 29.1 63.7 64.7 23.1 23.7 16.0 12.0 31.4 27.5 38.2 19.6
NSMT P.70407 G. c a u e r e n s i s female 29.5 Lombok 16.0 16.0 27.0 10.0 9.0 28.1 65.1 57.8 21.7 21.0 15.6 11.5 28.9 31.3 28.9 18.1
SAIAB 69164 G. c a u e r e n s i s female 36.5 Yemen, Red Sea 17.0 17.0 26.0 9.5 9.0 29.0 70.8 68.9 23.0 24.4 16.7 11.5 31.1 26.4 33.0 18.9
SAIAB 73710 G. c a u e r e n s i s male 41.5 Aliwal Shoal, South
Africa
17.0 17.0 30.0 9.0 9.0 27.5 69.3 64.0 21.2 20.5 17.3 11.3 34.2 26.3 30.7 21.1
SAIAB 73710 G. c a u e r e n s i s male 35.5 Aliwal Shoal, South
Africa
16.0 17.0 27.0 10.0 9.0 27.3 74.2 76.3 20.8 23.4 16.9 11.0 34.0 26.8 34.0 23.7
SAIAB 73710 G. c a u e r e n s i s female 41.0 Aliwal Shoal, South
Africa
18.0 17.0 27.0 10.0 10.0 28.5 64.1 68.4 22.0 23.2 16.6 12.0 31.6 24.8 31.6 20.5
SAIAB 73710 G. c a u e r e n s i s female 40.5 Aliwal Shoal, South
Africa
18.0 17.0 27.0 11.0 10.0 29.1 65.3 62.7 21.5 22.5 17.0 12.1 31.4 23.7 28.0 20.3
USNM 210157 G. c a u e r e n s i s male 31.5 Ceram, Moluccas 15.0 16.0 26.0 10.5 9.0 27.6 67.8 62.1 21.9 22.2 16.2 11.7 28.7 28.7 34.5 16.1
USNM 210157 G. c a u e r e n s i s female 30.0 Ceram, Moluccas 16.0 16.0 27.0 10.0 8.0 27.7 66.3 65.1 21.7 22.7 16.7 11.0 28.9 27.7 33.7 18.1
USNM 210157 G. c a u e r e n s i s female 27.0 Ceram, Moluccas 16.0 16.0 27.0 10.5 8.0 27.8 68.0 70.7 22.6 23.0 17.4 11.1 30.7 29.3 30.7 18.7
USNM 210157 G. c a u e r e n s i s female 28.0 Ceram, Moluccas 17.0 16.0 28.0 10.5 9.0 28.2 65.8 62.0 20.4 22.1 15.0 11.1 27.8 27.8 31.6 17.7
USNM 308212 G. c a u e r e n s i s male 39.0 One Tree Island, Qld 17.0 16.0 26.0 10.0 8.0 26.7 71.2 67.3 22.8 23.3 15.6 12.6 28.8 27.9 32.7 20.2
USNM 308212 G. c a u e r e n s i s male 34.0 One Tree Island, Qld 16.0 16.0 27.0 10.0 8.0 26.8 71.4 68.1 22.6 23.8 17.4 12.9 28.6 27.5 29.7 18.7
USNM 308212 G. c a u e r e n s i s male 39.5 One Tree Island, Qld 16.0 16.0 27.0 10.0 8.0 27.6 74.3 63.3 22.3 23.8 16.7 12.2 30.3 24.8 34.9 20.2
USNM 308212 G. c a u e r e n s i s female 34.5 One Tree Island, Qld 15.0 16.0 27.0 10.0 9.0 27.2 67.0 68.1 20.3 22.0 15.7 11.6 29.8 24.5 29.8 19.1
USNM 327456 G. c a u e r e n s i s male 48.0 Taiwan 16.0 16.0 27.0 9.5 9.0 25.8 74.2 72.6 23.1 23.1 17.3 12.5 33.1 24.2 36.3 21.0
USNM 327456 G. c a u e r e n s i s male 47.0 Taiwan 15.0 15.0 27.0 9.5 9.0 26.0 73.8 68.9 21.9 22.6 16.0 11.3 35.2 25.4 35.2 20.5
USNM 327456 G. c a u e r e n s i s female 44.5 Taiwan 15.0 15.0 26.0 10.0 8.0 26.5 66.9 73.7 21.8 21.6 16.4 11.2 33.1 25.4 34.7 25.4
USNM 327456 G. c a u e r e n s i s female 42.0 Taiwan 16.0 16.0 26.0 9.5 8.0 27.4 65.2 67.0 22.4 22.4 16.7 11.7 33.9 25.2 32.2 20.0
USNM 327456 G. c a u e r e n s i s female 42.5 Taiwan 16.0 17.0 28.0 10.0 9.0 26.1 69.4 68.5 22.6 23.5 17.9 11.3 32.4 25.2 33.3 22.5
USNM 379630 G. c a u e r e n s i s female 40.5 Rapa 16.0 16.0 27.0 10.0 9.0 27.4 69.4 63.1 21.7 22.2 17.3 11.9 37.8 26.1 35.1 22.5
USNM 379718 G. c a u e r e n s i s male 39.0 Rapa 15.0 16.0 26.0 9.5 10.0 26.9 70.5 61.9 22.3 22.1 15.6 11.8 34.3 27.6 35.2 21.0
USNM 379718 G. c a u e r e n s i s female 35.0 Rapa 14.0 14.0 29.0 10.0 8.0 28.9 74.3 62.4 24.3 24.6 15.1 12.0 34.7 27.7 32.7 23.8
USNM 379718 G. c a u e r e n s i s female 35.5 Rapa 17.0 17.0 27.0 11.0 9.0 28.2 72.0 71.0 21.4 22.3 17.7 10.4 35.0 29.0 34.0 22.0
USNM 379741 G. c a u e r e n s i s male 37.5 Rapa 16.0 16.0 27.0 10.5 9.0 26.7 64.0 60.0 21.9 21.6 15.5 12.3 32.0 28.0 35.0 18.0
USNM 379741 G. c a u e r e n s i s male 35.0 Rapa 17.0 17.0 28.0 10.5 9.0 26.9 73.4 58.5 20.9 22.3 15.7 11.4 35.1 27.7 38.3 22.3
USNM 379741 G. c a u e r e n s i s male 28.5 Rapa 16.0 17.0 28.0 10.0 10.0 27.7 64.6 60.8 18.9 20.7 17.5 11.2 31.6 29.1 34.2 20.3
...... continued on the next page
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.

Zootaxa 3529 © 2012 Magnolia Press · 67
A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
APPENDIX 1. (Continued)
Sample ID
(catalogue number
Species Sex SL (mm) Location
Counts and proportions used in CAP analysis
Pectoral right
Pectoral left
Lat.scales
TRB
Pred.scales
HL in SL
HD in HL
HW in HL
BDA in SL
BD D1 in SL
CPL in SL
CPD in SL
Snout in HL
Eye in HL
Jaw in HL
Preorb. in HL
USNM 379757 G. c a u e r e n s i s male 32.0 Rapa 16.0 16.0 28.0 9.0 8.0 26.3 66.7 67.9 18.8 19.7 16.6 11.6 31.0 27.4 35.7 20.2
USNM 379787 G. c a u e r e n s i s female 31.0 Rapa 16.0 16.0 28.0 9.0 8.0 28.7 59.6 62.9 20.3 20.0 16.5 11.6 38.2 30.3 32.6 22.5
USNM 379839 G. c a u e r e n s i s female 40.0 Anarua Bay, Rapa 18.0 18.0 28.0 10.0 9.0 28.0 70.5 75.0 20.0 20.5 16.8 11.3 38.4 27.7 35.7 23.2
USNM 379839 G. c a u e r e n s i s female 38.5 Anarua Bay, Rapa 17.0 17.0 27.0 11.0 8.0 27.3 73.3 72.4 22.1 22.1 17.7 11.9 38.1 25.7 34.3 23.8
BPBM 19665 G. k ni g h t i male 62.0 Oahu, Hawaii 16.0 16.0 27.0 10.5 10.0 25.0 81.9 77.4 21.9 22.6 16.5 12.6 43.2 27.1 38.1 24.5
BPBM 19665 G. k ni g h t i male 64.0 Oahu, Hawaii 16.0 16.0 27.0 11.5 8.0 26.9 75.6 68.6 23.4 24.7 15.2 12.5 34.9 25.0 34.9 23.3
BPBM 19665 G. k ni g h t i male 59.0 Oahu, Hawaii 16.0 17.0 28.0 11.5 10.0 27.1 85.6 73.8 24.6 23.7 16.9 12.7 37.5 26.3 40.6 23.8
BPBM 19665 G. k ni g h t i male 55.0 Oahu, Hawaii 16.0 16.0 28.0 11.0 9.0 27.6 81.6 75.7 24.0 25.1 16.2 13.1 39.5 27.6 36.2 23.0
BPBM 19665 G. k ni g h t i female 61.0 Oahu, Hawaii 16.0 16.0 27.0 10.5 7.0 27.9 74.1 65.3 24.6 25.6 18.7 12.5 36.5 27.1 37.1 24.7
BPBM 19665 G. k ni g h t i female 59.0 Oahu, Hawaii 16.0 15.0 27.0 10.0 9.0 27.6 71.8 69.9 25.4 25.4 16.8 11.9 35.6 27.6 34.4 21.5
BPBM 19665 G. k ni g h t i female 58.0 Oahu, Hawaii 17.0 16.0 28.0 11.0 8.0 27.8 75.2 67.1 24.5 25.0 16.7 12.4 34.2 26.7 36.6 22.4
BPBM 19665 G. k ni g h t i female 59.0 Oahu, Hawaii 17.0 16.0 27.0 11.0 8.0 28.0 76.4 80.0 24.6 24.9 15.9 12.4 40.6 24.8 37.0 27.9
BPBM 19665 G. k ni g h t i female 60.0 Oahu, Hawaii 17.0 17.0 26.0 11.5 8.0 28.3 73.5 72.9 26.0 24.7 16.7 13.5 41.2 27.1 34.1 24.1
BPBM 19665 G. k ni g h t i female 61.0 Oahu, Hawaii 17.0 16.0 27.0 11.0 8.0 27.2 72.3 68.7 25.7 24.3 16.6 12.8 38.0 25.3 33.7 24.7
BPBM 28720 G. k ni g h t i female 42.5 South Kahala, Hawaii 16.0 16.0 26.0 11.0 9.0 29.9 72.4 65.4 25.2 26.1 17.6 12.0 37.0 28.3 37.0 21.3
BPBM 34873 G. k ni g h t i male 36.0 Midway Atoll 16.0 17.0 26.0 10.0 9.0 26.9 70.1 68.0 21.1 22.5 17.5 11.4 29.9 32.0 36.1 23.7
BPBM 34873 G. k ni g h t i male 34.0 Midway Atoll 16.0 16.0 27.0 9.0 9.0 27.9 69.5 67.4 21.5 23.2 16.5 11.5 32.6 31.6 31.6 22.1
BPBM 34873 G. k ni g h t i female 35.5 Midway Atoll 17.0 17.0 27.0 10.0 8.0 27.6 66.3 61.2 20.6 23.4 18.3 10.7 28.6 30.6 36.7 20.4
CAS/SU 7468 G. kn i g h t i male 43.0 Paratype G. k n i g h t i 15.0 16.0 27.0 10.0 9.0 26.7 72.2 67.8 21.6 22.3 17.0 12.1 33.0 25.2 34.8 19.1
CAS/SU 7468 G. kn i g h t i male 43.0 Paratype G. k n i g h t i 16.0 16.0 27.0 11.0 9.0 26.7 76.5 78.3 23.3 23.3 17.2 11.4 35.7 26.1 38.3 19.1
CAS/SU 7468 G. kn i g h t i male 32.5 Paratype G. k n i g h t i 16.0 15.0 27.0 10.0 10.0 26.5 72.1 67.4 22.2 23.4 16.9 12.0 32.6 29.1 33.7 19.8
CAS/SU 7468 G. kn i g h t i female 38.0 Paratype G. k n i g h t i 15.0 16.0 27.0 10.0 8.0 26.3 75.0 76.0 21.1 22.4 18.9 10.5 44.0 27.0 33.0 23.0
CAS/SU 7468 G. kn i g h t i female 39.5 Paratype G. k n i g h t i 16.0 16.0 27.0 9.5 9.0 26.8 75.5 72.6 22.8 24.6 19.0 11.6 34.0 28.3 35.8 21.7
USNM 78072 G. k n ig h t i male 40.5 Hawaii 16.0 16.0 25.0 9.0 9.0 26.4 77.6 75.7 23.5 23.0 15.8 11.6 36.4 29.0 36.4 20.6
USNM 78072 G. k n ig h t i female 44.0 Hawaii 16.0 16.0 27.0 9.5 8.0 28.6 75.4 73.8 23.6 23.9 15.2 11.8 38.1 27.0 29.4 22.2
AMS I.21938–003 G. o p h th a l mo t a e n ia male 35.5 Cebu, Philippines 16.0 17.0 26.0 9.5 8.0 28.2 75.0 71.0 25.1 25.1 16.1 11.8 33.0 28.0 35.0 17.0
AMS I.21938–003 G. o p h th a l mo t a e n ia male 24.0 Cebu, Philippines 16.0 16.0 26.0 9.0 8.0 28.3 73.5 73.5 22.9 25.8 15.4 11.7 35.3 29.4 32.4 20.6
AMS I.21938–003 G. o p h th a l mo t a e n ia male 27.0 Cebu, Philippines 16.0 16.0 28.0 10.5 9.0 28.5 72.7 71.4 22.2 24.1 17.0 11.5 32.5 29.9 35.1 19.5
AMS I.21938–003 G. o p h th a l mo t a e n ia female 23.5 Cebu, Philippines 16.0 16.0 27.0 9.0 9.0 27.7 73.8 72.3 21.3 24.7 16.2 11.1 32.3 30.8 30.8 20.0
AMS I.24136–001 G. o p h th a l mo t a e n ia male 36.5 Batangas, Philippines 16.0 16.0 27.0 9.5 8.0 26.8 78.6 72.4 23.3 12.9 16.4 11.8 37.8 27.6 35.7 21.4
AMS I.31088–002 G. o p h th a l mo t a e n ia male 25.0 Solomon Islands, W
Pacific
16.0 16.0 25.0 10.0 8.0 29.2 68.5 61.6 23.2 11.2 16.0 11.6 31.5 28.8 35.6 20.5
AMS I.32492–009 G. o p h th a l mo t a e n ia male 26.0 Madang, New Guinea 15.0 17.0 25.0 9.0 8.0 28.1 75.3 69.9 25.4 25.0 16.5 12.7 34.2 31.5 32.9 20.5
BPBM 18670 G. o ph t h al m o t a en i a male 52.0 Neotype G. d a v ao e n si s 10.0 16.0 30.0 10.0 9.0 25.2 82.4 73.3 25.0 25.6 18.5 12.5 35.9 26.7 42.0 19.8
BPBM 38374 G. o ph t h al m o t a en i a male 51.0 Taiwan 16.0 16.0 26.0 10.5 10.0 26.7 79.4 71.3 26.9 25.1 18.6 13.1 34.6 25.7 40.4 19.9
BPBM 38374 G. o ph t h al m o t a en i a male 52.0 Taiwan 17.0 16.0 26.0 10.5 8.0 26.2 86.0 66.2 26.3 25.2 15.6 14.0 33.1 25.7 40.4 22.1
CAS 51506 G. o ph t h al m o t a en i a male 34.0 Solomon Islands, W
Pacific
16.0 16.0 25.0 9.0 8.0 25.9 79.5 72.7 23.8 24.4 16.2 12.1 34.1 29.5 38.6 21.6
CAS 51506 G. o ph t h al m o t a en i a male 33.5 Solomon Islands, W
Pacific
16.0 16.0 24.0 9.5 8.0 26.9 76.7 68.9 23.9 24.2 16.7 11.9 30.0 30.0 35.6 16.7
CAS 51506 G. o ph t h al m o t a en i a male 33.5 Solomon Islands, W
Pacific
15.0 15.0 25.0 9.0 8.0 26.9 78.9 67.8 25.4 25.4 17.3 12.5 31.1 28.9 36.7 20.0
CAS 51506 G. o ph t h al m o t a en i a male 32.5 Solomon Islands, W
Pacific
15.0 16.0 26.0 9.5 8.0 27.1 73.9 69.3 24.3 25.2 16.0 12.6 28.4 29.5 34.1 19.3
CAS 51506 G. o ph t h al m o t a en i a male 32.5 Solomon Islands, W
Pacific
16.0 16.0 26.0 10.5 7.0 26.8 75.9 66.7 24.3 24.9 17.5 12.3 31.0 28.7 34.5 20.7
CAS 51506 G. o ph t h al m o t a en i a male 34.5 Solomon Islands, W
Pacific
16.0 16.0 27.0 9.5 9.0 26.4 78.0 71.4 23.8 25.5 17.1 12.2 25.3 28.6 33.0 17.6
CAS 51506 G. o ph t h al m o t a en i a male 33.0 Solomon Islands, W
Pacific
16.0 15.0 26.0 10.5 8.0 27.3 77.8 70.0 24.2 24.8 17.3 12.1 30.0 27.8 35.6 20.0
...... continued on the next page
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LARSON & BUCKLE
68 · Zootaxa 3529 © 2012 Magnolia Press
APPENDIX 1. (Continued)
Sample ID
(catalogue number
Species Sex SL (mm) Location
Counts and proportions used in CAP analysis
Pectoral right
Pectoral left
Lat.scales
TRB
Pred.scales
HL in SL
HD in HL
HW in HL
BDA in SL
BD D1 in SL
CPL in SL
CPD in SL
Snout in HL
Eye in HL
Jaw in HL
Preorb. in HL
CAS 51506 G. o ph t h al m o t a en i a male 31.5 Solomon Islands, W
Pacific
16.0 16.0 25.0 9.5 8.0 27.6 77.0 69.0 23.8 25.1 16.5 12.4 32.2 29.9 34.5 19.5
CAS 51506 G. o ph t h al m o t a en i a female 30.5 Solomon Islands, W
Pacific
16.0 16.0 25.0 10.0 8.0 28.2 73.3 66.3 24.6 27.5 17.4 11.5 30.2 29.1 32.6 17.4
CAS 51506 G. o ph t h al m o t a en i a female 30.0 Solomon Islands, W
Pacific
16.0 16.0 26.0 9.5 9.0 28.3 77.6 67.1 25.3 27.0 16.7 12.0 29.4 29.4 34.1 17.6
CAS 51506 G. o ph t h al m o t a en i a female 30.0 Solomon Islands, W
Pacific
16.0 16.0 26.0 9.5 8.0 27.7 79.5 74.7 24.3 26.7 16.7 12.0 30.1 31.3 37.3 21.7
CAS 51506 G. o ph t h al m o t a en i a female 31.0 Solomon Islands, W
Pacific
16.0 16.0 25.0 10.0 8.0 27.4 77.6 65.9 24.5 26.5 19.4 11.9 31.8 29.4 34.1 18.8
MSMT–P.67349 G. o p h t ha l m o t ae n i a male 33.0 Iriomote–jima, S Japan 16.0 16.0 26.0 10.0 9.0 27.0 76.4 68.5 24.2 24.8 18.2 12.7 32.6 30.3 37.1 19.1
NSMT–P.64505 G. o p h t ha l m o ta e n ia male 26.5 Ceram, Indonesia 16.0 16.0 27.0 10.5 8.0 28.3 70.7 68.0 22.3 23.0 15.8 11.7 33.3 26.7 32.0 18.7
NSMT–P.64545 G. o p h t ha l m o ta e n ia male 22.5 Ambon, Indonesia 16.0 15.0 27.0 9.5 8.0 30.7 72.5 62.3 22.7 24.4 16.9 11.6 33.3 29.0 29.0 18.8
NSMT–P.64545 G. o p h t ha l m o ta e n ia female 27.0 Ambon, Indonesia 16.0 16.0 26.0 10.0 8.0 30.0 79.0 74.1 24.1 25.2 15.6 12.2 37.0 27.2 37.0 22.2
NSMT–P.64639 G. o p h t ha l m o ta e n ia female 27.0 Ceram, Indonesia 16.0 15.0 27.0 9.0 8.0 29.6 72.5 63.8 24.4 26.3 16.3 12.2 33.8 32.5 32.5 22.5
NTM S.11328–002 G. o p ht h a lm o t a e ni a male 34.0 Solomon Islands, W
Pacific
17.0 16.0 27.0 9.0 9.0 26.8 79.1 72.5 24.4 12.4 17.1 11.2 35.2 30.8 35.2 19.8
NTM S.12697–001 G. o p ht h a lm o t a e ni a male 29.0 Poso, Sulawesi,
Indonesia
17.0 17.0 27.0 9.5 8.0 29.0 75.0 71.4 24.8 25.9 17.9 12.8 36.9 27.4 33.3 20.2
NTM S.13679–031 G. o p ht h a lm o t a e ni a female 33.5 Madang, New Guinea 16.0 17.0 25.0 10.0 6.0 28.7 82.3 76.0 25.1 26.9 14.6 12.5 36.5 26.0 34.4 24.0
USNM 266393 G. o p ht h a lm o t a en i a female 35.5 Halmahera, Indonesia 16.0 16.0 26.0 9.0 7.0 28.7 70.6 57.8 22.5 23.9 16.1 11.0 28.4 26.5 33.3 19.6
USNM 308210 G. o p ht h a lm o t a en i a male 34.0 Milne Bay, New Guinea 16.0 16.0 27.0 10.5 9.0 27.4 74.2 69.9 24.4 24.1 15.6 11.5 30.1 31.2 34.4 19.4
USNM 308210 G. o p ht h a lm o t a en i a male 27.0 Milne Bay, New Guinea 15.0 16.0 27.0 9.5 9.0 28.9 73.1 70.5 23.0 25.9 16.7 11.9 32.1 29.5 34.6 19.2
USNM 308210 G. o p ht h a lm o t a en i a male 25.0 Milne Bay, New Guinea 16.0 16.0 26.0 10.0 8.0 28.8 70.8 69.4 22.8 24.4 17.2 11.2 33.3 29.2 33.3 20.8
USNM 308210 G. o p ht h a lm o t a en i a male 30.0 Milne Bay, New Guinea 16.0 16.0 27.0 10.5 8.0 28.3 74.1 70.6 24.3 26.0 16.0 12.3 35.3 30.6 36.5 20.0
USNM 308210 G. o p ht h a lm o t a en i a male 28.0 Milne Bay, New Guinea 16.0 16.0 26.0 9.0 8.0 28.2 75.9 68.4 23.6 25.7 16.4 11.8 32.9 30.4 32.9 20.3
USNM 308210 G. o p ht h a lm o t a en i a female 27.0 Milne Bay, New Guinea 16.0 16.0 25.0 10.5 8.0 29.3 74.7 70.9 24.4 26.3 16.7 11.9 31.6 27.8 32.9 20.3
USNM 308210 G. o p ht h a lm o t a en i a female 27.5 Milne Bay, New Guinea 15.0 16.0 27.0 9.5 9.0 29.1 70.0 71.3 23.6 25.5 16.0 11.6 31.3 30.0 30.0 18.8
USNM 308210 G. o p ht h a lm o t a en i a female 24.0 Milne Bay, New Guinea 16.0 16.0 26.0 9.5 8.0 28.8 73.9 72.5 22.9 25.4 15.8 11.7 30.4 30.4 30.4 21.7
USNM 308210 G. o p ht h a lm o t a en i a female 23.0 Milne Bay, New Guinea 16.0 16.0 26.0 10.5 9.0 29.1 68.7 67.2 22.2 24.3 15.7 11.7 26.9 29.9 29.9 19.4
USNM 308210 G. o p ht h a lm o t a en i a female 19.5 Milne Bay, New Guinea 16.0 16.0 25.0 9.5 8.0 29.2 71.9 71.9 22.6 25.6 15.9 11.3 28.1 31.6 28.1 17.5
USNM 308210 G. o p ht h a lm o t a en i a female 22.0 Milne Bay, New Guinea 16.0 16.0 26.0 9.0 8.0 28.2 74.2 71.0 20.9 24.5 15.9 10.9 29.0 33.9 32.3 19.4
BPBM 32850 G. p a s cu e n si s female 34.0 Holotype G. c a u e r e n s i s
pascuensis
19.0 19.0 27.0 10.0 9.0 26.8 65.9 63.7 23.2 22.9 16.2 11.8 30.8 30.8 36.3 18.7
BPBM 32851 G. p a s cu e n si s male 39.0 Easter Island 19.0 19.0 27.0 10.0 8.0 29.0 70.8 75.2 21.8 21.8 16.4 13.1 39.8 26.5 35.4 23.0
BPBM 32851 G. p a s cu e n si s female 35.0 Easter Island 18.0 18.0 27.0 10.0 9.0 30.3 66.0 70.8 21.1 22.9 15.7 12.0 34.9 28.3 32.1 19.8
BPBM 32852 G. p a s cu e n si s female 41.5 Easter Island 19.0 19.0 27.0 9.5 10.0 29.4 63.9 67.2 21.9 21.7 15.7 11.3 35.2 27.0 34.4 19.7
BPBM 32853 G. p a s cu e n si s female 45.0 Easter Island 18.0 18.0 27.0 9.5 8.0 27.8 68.0 70.4 22.4 23.3 14.4 11.8 36.8 28.0 32.8 24.0
LACM 5974 G. t h om p s on i male 58.0 Jamaica 17.0 17.0 28.0 10.5 10.0 26.4 68.6 74.5 22.2 21.9 16.7 12.2 35.3 24.2 34.6 19.6
LACM 5974 G. t h om p s on i male 53.0 Jamaica 17.0 17.0 27.0 10.5 10.0 25.7 73.5 77.2 22.1 21.1 17.2 12.1 38.2 25.0 39.0 22.8
LACM 5974 G. t h om p s on i male 35.5 Jamaica 17.0 17.0 27.0 10.0 11.0 27.6 69.4 66.3 23.1 24.2 16.9 11.8 32.7 25.5 34.7 20.4
LACM 5974 G. t h om p s on i male 49.0 Jamaica 16.0 16.0 26.0 10.5 10.0 26.5 71.5 71.5 22.7 22.7 16.7 12.0 32.3 23.1 36.2 20.0
LACM 5974 G. t h om p s on i male 45.0 Jamaica 17.0 17.0 27.0 10.0 10.0 27.1 69.7 69.7 23.3 23.6 17.8 12.9 32.8 24.6 37.7 20.5
LACM 5974 G. t h om p s on i male 40.5 Jamaica 17.0 17.0 27.0 10.0 10.0 26.4 69.2 72.9 22.0 23.0 15.8 11.9 29.9 24.3 36.4 20.6
LACM 5974 G. t h om p s on i male 36.5 Jamaica 17.0 17.0 26.0 10.5 9.0 27.7 64.4 67.3 22.7 22.5 15.6 12.3 29.7 24.8 35.6 17.8
LACM 5974 G. t h om p s on i female 46.0 Jamaica 17.0 17.0 27.0 10.0 11.0 26.7 69.9 69.9 22.6 23.5 17.8 12.4 32.5 24.4 35.8 20.3
LACM 5974 G. t h om p s on i female 44.0 Jamaica 17.0 17.0 27.0 10.5 10.0 29.1 71.1 70.3 25.7 25.2 14.3 13.2 32.8 24.2 35.2 20.3
LACM 5974 G. t h om p s on i female 35.0 Jamaica 16.0 16.0 27.0 9.0 10.0 28.0 69.4 71.4 24.0 23.7 16.6 12.0 28.6 25.5 32.7 20.4
LACM 5974 G. t h om p s on i female 41.5 Jamaica 17.0 17.0 27.0 10.5 10.0 28.0 65.5 69.8 23.1 22.7 17.1 12.0 31.9 24.1 32.8 21.6
LACM 5974 G. t h om p s on i female 34.0 Jamaica 17.0 17.0 27.0 10.0 9.0 28.2 67.7 67.7 22.6 23.5 14.7 11.8 30.2 26.0 32.3 17.7
USNM 179252 G. t ho m p so n i male 51.0 Ven z u el a 16.0 17.0 28.0 10.5 10.0 27.1 68.8 68.8 22.4 21.6 16.5 12.2 34.1 23.2 36.2 21.7
...... continued on the next page
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Zootaxa 3529 © 2012 Magnolia Press · 69
A REVISION OF THE GOBY GENUS GNATHOLEPIS BLEEKER
APPENDIX 1. (Continued)
Sample ID
(catalogue number
Species Sex SL (mm) Location
Counts and proportions used in CAP analysis
Pectoral right
Pectoral left
Lat.scales
TRB
Pred.scales
HL in SL
HD in HL
HW in HL
BDA in SL
BD D1 in SL
CPL in SL
CPD in SL
Snout in HL
Eye in HL
Jaw in HL
Preorb. in HL
USNM 179252 G. t ho m p so n i male 45.0 Ven z u el a 18.0 17.0 28.0 10.0 9.0 26.2 69.5 77.1 19.8 21.3 17.1 11.1 32.2 28.0 36.4 19.5
USNM 179252 G. t ho m p so n i male 38.5 Ven z u el a 18.0 17.0 28.0 10.0 10.0 27.8 71.0 65.4 20.8 22.3 17.4 11.9 28.0 29.0 35.5 18.7
USNM 179252 G. t ho m p so n i female 46.0 Ven z u el a 17.0 17.0 26.0 11.0 9.0 27.8 64.8 67.2 22.6 23.0 16.3 11.5 30.5 28.1 28.1 20.3
USNM 179252 G. t ho m p so n i female 42.0 Ven z u el a 17.0 17.0 27.0 10.0 10.0 27.1 67.5 68.4 21.0 21.4 15.5 11.0 33.3 28.1 36.8 20.2
USNM 179253 G. t ho m p so n i male 39.0 Ven e z u el a 16.0 17.0 26.0 10.5 10.0 27.2 64.2 67.9 21.5 21.3 15.9 12.1 28.3 27.4 34.9 17.9
USNM 179254 G. t ho m p so n i male 32.0 Ven e z u el a 17.0 17.0 27.0 10.0 11.0 28.4 69.2 65.9 21.9 23.1 16.6 11.6 28.6 26.4 36.3 19.8
USNM 179254 G. t ho m p so n i male 37.0 Ven e z u el a 17.0 16.0 26.0 10.0 10.0 27.0 67.0 69.0 20.5 20.8 15.1 11.9 31.0 27.0 36.0 20.0
USNM 218840 G. t ho m p so n i male 44.0 Ascension Island 18.0 18.0 27.0 11.5 10.0 27.0 69.7 65.5 17.7 22.0 15.9 11.8 29.4 26.1 36.1 16.8
USNM 264886 G. t ho m p so n i male 58.0 Barbuda 18.0 18.0 28.0 10.5 9.0 27.6 71.9 67.5 24.3 24.3 17.2 13.8 31.3 23.8 35.0 21.9
USNM 267896 G. t ho m p so n i male 29.0 St Helena 16.0 16.0 28.0 10.0 9.0 28.3 58.5 57.3 21.4 19.3 16.6 10.3 28.0 28.0 32.9 17.1
USNM 276131 G. t ho m p so n i female 26.5 Belize 17.0 17.0 26.0 11.0 10.0 27.5 67.1 65.8 21.9 23.4 17.0 11.3 27.4 30.1 31.5 17.8
USNM 276135 G. t ho m p so n i male 41.5 Belize 17.0 17.0 26.0 11.5 11.0 27.7 71.3 71.3 21.0 21.9 14.7 11.6 30.4 27.0 34.8 20.9
USNM 276135 G. t ho m p so n i male 36.5 Belize 16.0 16.0 26.0 10.0 11.0 27.4 64.0 60.0 20.3 19.7 17.0 11.5 29.0 32.0 32.0 21.0
USNM 276135 G. t ho m p so n i male 38.0 Belize 18.0 17.0 27.0 11.0 10.0 27.1 69.9 68.0 21.6 21.3 15.5 12.1 34.0 26.2 38.8 21.4
USNM 276135 G. t ho m p so n i male 27.5 Belize 18.0 17.0 27.0 9.5 9.0 28.4 67.9 66.7 20.7 21.8 17.5 11.3 26.9 28.2 34.6 17.9
USNM 276135 G. t ho m p so n i female 27.0 Belize 17.0 16.0 27.0 11.0 9.0 29.6 63.8 61.3 21.5 21.9 16.7 11.9 28.8 27.5 33.8 22.5
USNM 317097 G. t ho m p so n i male 41.0 Tobago 17.0 17.0 27.0 10.0 10.0 27.6 62.8 63.7 20.2 21.7 16.3 11.0 29.2 27.4 37.2 17.7
USNM 317097 G. t ho m p so n i male 33.5 Tobago 17.0 17.0 27.0 10.5 9.0 28.4 63.2 64.2 21.5 23.0 16.1 11.6 28.4 26.3 35.8 16.8
USNM 317097 G. t ho m p so n i male 32.0 Tobago 17.0 17.0 28.0 11.0 9.0 28.1 61.1 61.1 20.9 20.6 16.3 11.3 28.9 27.8 33.3 16.7
USNM 317097 G. t ho m p so n i female 33.5 Tobago 17.0 17.0 26.0 10.0 10.0 29.9 62.0 60.0 21.8 23.9 16.4 11.6 30.0 27.0 31.0 19.0
USNM 317097 G. t ho m p so n i female 38.0 Tobago 17.0 16.0 27.0 11.0 11.0 28.7 61.5 65.1 21.6 23.7 16.6 11.3 30.3 25.7 33.0 19.3
USNM 317098 G. t ho m p so n i male 31.5 Tobago 17.0 17.0 29.0 10.5 11.5 27.3 64.0 62.8 19.7 21.3 17.5 10.8 31.4 27.9 34.9 17.4
USNM 317098 G. t ho m p so n i male 34.0 Tobago 16.0 16.0 27.0 11.0 10.0 27.4 64.5 68.8 19.7 21.2 15.6 11.2 29.0 26.9 34.4 18.3
USNM 317098 G. t ho m p so n i female 21.5 Tobago 18.0 18.0 26.0 10.0 10.0 29.8 64.1 59.4 20.0 21.9 18.1 10.2 29.7 29.7 32.8 18.8
USNM 320763 G. t ho m p so n i male 36.5 Nicaragua 17.0 18.0 28.0 10.0 10.0 27.1 74.7 74.7 20.0 20.8 14.8 12.3 37.4 28.3 36.4 20.2
USNM 346472 G. t ho m p so n i male 33.5 Belize 17.0 17.0 26.0 10.0 9.0 28.1 76.6 75.5 20.9 22.4 15.8 11.6 31.9 27.7 37.2 19.1
USNM 346472 G. t ho m p so n i male 27.0 Belize 16.0 16.0 28.0 10.0 9.0 29.6 68.8 75.0 21.1 21.5 17.0 11.1 33.8 27.5 32.5 20.0
USNM 357711 G. t h o m p so n i male 34.5 Brazil 16.0 17.0 26.0 10.5 10.0 27.2 67.0 69.1 22.3 22.6 15.9 12.5 28.7 26.6 36.2 21.3
USNM 357711 G. t h o m p so n i male 39.0 Brazil 16.0 16.0 26.0 10.5 9.0 28.2 63.6 70.0 21.5 23.1 17.4 12.1 30.9 26.4 34.5 13.6
USNM 357711 G. t h o m p so n i female 40.5 Brazil 16.0 16.0 27.0 10.0 9.0 28.6 66.4 71.6 22.5 23.5 16.5 12.3 34.5 25.9 34.5 20.7
USNM 362225 G. t ho m p so n i male 30.0 Nicaragua 17.0 17.0 28.0 9.0 10.0 29.0 72.4 72.4 20.7 23.7 16.7 12.3 36.8 27.6 34.5 23.0
USNM 362225 G. t ho m p so n i male 25.0 Nicaragua 18.0 18.0 26.0 10.0 10.0 30.0 65.3 69.3 18.0 21.6 16.8 10.4 30.7 28.0 32.0 18.7
USNM 362225 G. t ho m p so n i female 24.5 Nicaragua 17.0 16.0 28.0 10.0 9.0 29.4 65.3 68.1 20.0 23.3 16.3 11.0 30.6 29.2 33.3 16.7
USNM 82520 G. t h o mp s o n i female 43.0 Cuba 17.0 17.0 26.0 11.0 10.0 28.6 71.5 70.7 24.4 25.3 15.1 12.1 34.1 26.8 35.8 22.8
USNM 82520 G. t h o mp s o n i female 38.0 Cuba 17.0 17.0 28.0 10.0 10.0 28.9 70.9 65.5 22.6 23.9 14.5 12.4 31.8 35.5 33.6 21.8
USNM 82520 G. t h o mp s o n i female 44.0 Cuba 18.0 17.0 28.0 12.0 11.0 28.4 69.6 63.2 23.2 23.4 16.6 12.0 35.2 25.6 34.4 24.8
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