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Revision of the fish family Kyphosidae (Teleostei: Perciformes)

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A molecular phylogenetic analysis with complete species sampling of the family Kyphosidae revealed several discrepancies with the current taxonomy. We thus undertook a complete taxonomic revision of all kyphosid genera, i.e. Kyphosus Lacepède, 1801, and the monotypic Hermosilla Jenkins and Evermann, 1889, Sectator Jordan and Evermann, 1903 and Neoscorpis Smith, 1931. Species delimitation was determined on the basis of congruence between (a) monophyletic groupings in the molecular phylogeny, and (b) clusters of morphological variation in type material. Twelve species are supported and redescribed. Both Hermosilla and Sectator are considered junior synonyms of Kyphosus. Kyphosus azureus (Jenkins & Evermann, 1889) and K. ocyurus (Jordan & Gilbert, 1882) are redescribed accordingly. We designate a neotype for Kyphosus cornelii (Whitley, 1944), as the original material is lost, and new material was collected at the type locality for this study to facilitate comparison with other species of Kyphosus. Kyphosus sandwicensis (sensu Sauvage, 1880) was found to be a junior synonym of K. elegans (Peters, 1869). Kyphosus incisor (Cuvier in Cuvier & Valenciennes, 1831) and K. analogus (Gill, 1862) are considered junior synonyms of K. vaigiensis (Quoy & Gaimard, 1825). Kyphosus gallveii (Cunningham, 1910), K. pacificus Sakai and Nakabo, 2004 and K. lutescens (Jordan & Gilbert, 1882) are all considered junior synonyms of K. sectatrix (Linnaeus, 1758). One of the two syntype specimens of K. sectatrix was identified as the holotype of Pimelepterus bosquii (Lacepède, 1802), and proved to be a specimen of K. bigibbus Lacepède, 1801. This specimen is re-assigned as a non-type of K. bigibbus. Full re-descriptions of the following valid species are presented: K. bigibbus, K. cinerascens (Forsskål, 1775), K. cornelii, K. elegans, K. hawaiiensis Sakai and Nakabo, 2004, K. gladius Knudsen and Clements, 2013, K. sydneyanus (Günther, 1886) and K. vaigiensis, together with a key to the family. The distribution of Kyphosus species is reconsidered based on our taxonomic revision, indicating that four species (K. bigibbus, K. cinerascens, K. sectatrix and K. vaigiensis) occur in both the Atlantic and Indo-Pacific regions.
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Accepted by Eric Hilton: 11 Oct. 2013; published: 24 Dec. 2013
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ZOOTAXA
Revision of the fish family Kyphosidae (Teleostei: Perciformes)
STEEN WILHELM KNUDSEN & KENDALL D. CLEMENTS
School of Biological Sciences, University of Auckland, Auckland, New Zealand
(e-mail: sknu003@aucklanduni.ac.nz)
Phone: (+64 9) 373 7599 ext. 88483, Fax.: (+64 9) 373 7417
Corresponding author: Steen Wilhelm Knudsen
Magnolia Press
Auckland, New Zealand
3751
KNUDSEN & CLEMENTS
2 · Zootaxa 3751 (1) © 2013 Magnolia Press
STEEN WILHELM KNUDSEN & KENDALL D. CLEMENTS
Revision of the fish family Kyphosidae (Teleostei: Perciformes)
(Zo
otaxa 3751)
101 pp.; 30 cm.
24 Dec. 2013
ISBN 978-1-77557-316-6 (paperback)
ISBN 978-1-77557-317-3 (Online edition)
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ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
Zootaxa 3751 (1) © 2013 Magnolia Press · 3
TAXONOMIC REVISION OF KYPHOSIDAE
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Genus Kyphosus Lacepède . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Genus Neoscorpis Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Key to the species of Kyphosidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Regional keys to the species of Kyphosidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Key to the Atlantic species of Kyphosidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Key to the Indo-Pacific species of Kyphosidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Key to the East Pacific species of Kyphosidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Kyphosus azureus (Jenkins & Evermann, 1889) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Kyphosus bigibbus Lacepède, 1801 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Kyphosus cinerascens (Forsskål, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Kyphosus cornelii (Whitley, 1944) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Kyphosus elegans (Peters, 1869) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Kyphosus gladius Kn
udsen & Clements, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Kyphosus hawaiiensis Sakai and Nakabo, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Kyphosus ocyurus (Jord
an & Gilbert, 1882) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Kyphosus sectatrix (L
innaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Kyphosus sydneyanus (Günther, 1886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Kyphosus vaigiensis (Quoy & Gaimard, 1825) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Comparative material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Dichistiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Girellidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Microcanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Scorpididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Abstract
A molecular phylogenetic analysis with complete species sampling of the family Kyphosidae revealed several discrepan-
cies with the current taxonomy. We thus undertook a complete taxonomic revision of all kyphosid genera, i.e. K
yphosus
Lacepède, 1801, and the monotypic Hermosilla Jenkins and Evermann, 1889, Sectator Jordan and Evermann, 1903 and
Neoscorpis Smith, 1931. Species delimitation was determined on the basis of congruence between (a) monophyletic
groupings in the molecular phylogeny, and (b) clusters of morphological variation in type material. Twelve species are
supported and redescribed. Both Hermosilla and Sectator are considered junior synonyms of Kyphosus. Kyphosus azureus
(Jenkins & Evermann, 1889) and K. ocyurus (Jordan & Gilbert, 1882) are redescribed accordingly. We designate a neotype
for Kyphosus cornelii (Whitley, 1944), as the original material is lost, and new material was collected at the type locality
for this study to facilitate comparison with other species of Kyphosus. Kyphosus sandwicensis (sensu Sauvage, 1880) was
found to be a junior synonym of K. elegans (Peters, 1869
). Kyphosus incisor (Cuvier in Cuvier & Valenciennes, 1831) and
K. analogus (Gill, 1862) are considered junior synonyms of K. vaigiensis (Quoy & Gaimard, 1825). Kyphosus gallveii (Cun-
ningham, 1910), K.
pacificus Sakai and Nakabo, 2004 and K. lutescens (Jordan & Gilbert, 1882) are all considered junior syn-
onyms of K. sectatr
ix (Linnaeus, 1758). One of the two syntype specimens of K. sectatrix was identified as the holotype of
Pimelepterus bosquii (Lacepède, 1802), and proved to be a specimen of K. bigibbus Lacepède, 1801. This specimen is re-as-
signed as a non-type of K. big
ibbus. Full re-descriptions of the following valid species are presented: K. bigibbus, K. cinerascens
(Forsskål, 1775), K. cornelii, K. elegans, K. hawaiiensis Sakai and Nakabo, 2004, K. gladius Knudsen and Clements, 2013, K.
sydneyanus (Günther, 1886) and K. vaigiensis, together with a key to the family. The distribution of Kyphosus species is recon-
sidered based on our taxonomic revision, indicating that four species (K. b
igibbus, K. cinerascens, K. sectatrix and K. vaigiensis)
occur in both the Atlantic and Indo-Pacific regions.
Key words: drummers, Girellidae, Hermosilla, Kyphosinae, Kyphosus, Neoscorpis, Rudderfishes, sea chubs, Sectator
KNUDSEN & CLEMENTS
4 · Zootaxa 3751 (1) © 2013 Magnolia Press
Introduction
The perciform family Kyphosidae currently contains 16 species in four genera: Hermosilla Jenkins and Evermann,
1889, Kyphosus Lacepède, 1801, Neoscorpis Smith, 1931 and Sectator Jordan and Evermann, 1903 (Randall 1968,
2005, Sakai & Nakabo 1995, 2004, 2006, Myers 1999, Kuiter & Debelius 2006, Knudsen & Clements 2013).
Hermosilla, Neoscorpis and Sectator are all monotypic, while Kyphosus currently consists of 13 species
(Eschmeyer 2012), including the recently described K. gladius (Knudsen & Clements 2013). All species of
Kyphosidae are herbivorous except for Sectator ocyurus, which feeds on zooplankton (Allen & Robertson 1997).
Kyphosids are moderate to large in size and occur in shallow waters (0–10 m depth).
They are associated with rocky shores and reefs worldwide at low latitudes, and at higher latitudes in the
M
editerranean Sea and the Pacific and Indian Oceans (Nelson 2006).
Of the four kyphosid genera, Hermosil
la is restricted to the central eastern Pacific (Allen & Robertson 1994,
Thomson et al. 2001). Kyphosus is widely distributed in the Atlantic, Indian and the Pacific Oceans (Randall 1968,
2005, 2007, Myers 1989, 1999, Kuiter & Debelius 2006). Kyphosus species occur in subtropical waters in both the
Southern and Northern hemispheres (Francis 1993, Clements 1997, Kuiter 1997, Last et al. 2011), where they have
been recorded from the Mediterranean (Dawson 1963, Merella et al. 1998, Francour & Mouine 2008, Relini et al.
2010, Ligas et al. 2011, Kiparissis et al. 2012), in the Red Sea (Randall 1983), and from temperate Australia (Allen
& Swainston 1988, Kuiter 1997, 2000, Randall et al. 1997) and northern New Zealand (Francis 1991, 1993, 1996,
Clements 1997). Markevich (2005) also reported the capture of Kyphosus in Peter the Great Bay, Russia.
Furthermore, Kyphosus is found in the central eastern (Dooley et al. 1985, Tortonese 1986, Santos et al. 1997,
Canas et al. 2005) and western Atlantic (Carvalho 1950, Randall 1968, Edwards & Glass 1987, Smith-Vaniz et al.
1999), in the Pacific Ocean from the central western part of the Pacific to Hawaii, Easter Island and the East Pacific
(Araga 1984, Allen & Robertson 1994, De la Cruz Agüero et al. 1997, Randall 2005, 2007, Kuiter & Debelius
2006, Randall & Cea 2010). The third genus, Sectator, is found across the Pacific from Central America across to
Japan (Araga 1984, Myers 1999, Randall 2007). The fourth and last genus, Neoscorpis, is restricted to eastern
South Africa and southern Mozambique and Fort Dauphin in Madagascar (Joubert & Hanekorn 1980, van der Elst
& King 2006).
The first record of the family Kyphosidae dates back to Linnaeus (1758), who described P
erca saltatrix and
later (Linnaeus 1766) corrected this to Perca sectatrix (currently valid as Kyphosus sectatrix). In 1775, Forsskål
described the species Sciæna cinerascens, tahmel from the Red Sea. The valid name for ‘Sciæna cinerascens,
tahmel’ is Kyphosus cinerascens, as the third name: ‘tahmel’ is a subspecific name that should not be used (see
Sakai & Nakabo 2006). Unfortunately, this specimen has been lost. Later, Lacepède (1801) described Kyphosus
bigibbus by monotypy. As the name Sciaena pertains to croakers (f. Sciaenidae) (Linnaeus 1758), the type species
designating the genus is Kyphosus bigibbus (Lacepède 1801). Additionally, Lacepède (1802) described
Pimelepterus (occasionally misspelled: Pimelipterus and Pimelopterus) and Xyster (Lacepède 1803), both later
considered junior synonyms of Kyphosus by Desoutter (1973). Whitley (1931) erected the subgenera
Leptokyphosus and Segutilum, both later designated as synonyms of Kyphosus (Scott 1971, Sakai & Nakabo 2006,
Knudsen & Clements 2013).
The higher taxonomic status of kyphosids is controversial. They were considered as the subfamily Kyphosinae
by
Nelson (2006) (i.e. comprising Hermosilla, Kyphosus, Neoscorpis and Sectator), who considered Kyphosidae to
comprise the subfamilies Girellinae, Kyphosinae, Microcanthinae, Parascorpidinae and Scorpidinae, i.e. the genera
Atypichthys, Bathystethus, Girella, Graus, Hermosilla, Kyphosus, Labracoglossa, Medialuna, Microcanthus,
Neatypus, Neoscorpis, Parascorpis, Scorpis, Sectator and Tilodon. However, an alternative approach has been to
assign each of these taxa family status (i.e. Girellidae, Kyphosidae, Microcanthidae, Parascorpididae and
Scorpididae) (Smith 1935, Horn 1984, Leis & van der Lingen 1997, Francis 2001, Randall 2005, Allen & Erdmann
2012), in which case the Kyphosidae comprises the genera Hermosilla, Kyphosus, Neoscorpis and Sectator
(Randall 2005). A subdivision of the family Pimelepteroidæ (which would equal Kyphosidae sensu Nelson 2006)
was put forward by Gill (1862), who regarded Pimelepteroidæ to contain the subfamilies Pimelepterinæ and
Girellinæ, with Pimelepterus in Pimelepterinæ and Girella in Girellinæ (Gill 1862). Johnson and Fritzsche (1989)
regarded Girellidae, Kyphosidae and Scorpididae as distinct, and pointed out, as did Johnson (1984), that the larval
form supports a close relationship between these taxa. Additionally, Freihofer (1963) found that the same pattern
10 of the ramus lateralis accesorius (RLA-10) supported a close relationship between Girellidae, Kyphosidae and
Zootaxa 3751 (1) © 2013 Magnolia Press · 5
TAXONOMIC REVISION OF KYPHOSIDAE
Scorpididae. Conversely, Neira et al. (1997) was unable to find support for this relationship in larval characters.
Molecular studies have shown that Oplegnathidae, Kuhliidae and Terapontidae fall within the clade containing
Girella, Kyphosus, Microcanthus and Scorpis (Yagishita et al. 2002, 2009, Near et al. 2012, 2013), and following
these molecular phylogenies we consider the family Kyphosidae sensu Randall (2005) (i.e. Kyphosinae sensu
Nelson 2006) to comprise the four genera Hermosilla, Neoscorpis, Kyphosus and Sectator, and unless specified
otherwise, all subsequent use of Kyphosidae follows this definition.
Although Horn (1984) pointed out a list of differences between Girellidae, Kyphosidae and Scorpididae (i
.e.
rows of premaxillary teeth and dentary teeth, branchiostegal rays, ray base preceeding procurrent spur, predorsal
bones, juvenile pigmentation and juvenile association), a full taxonomic designation of Kyphosidae is currently not
possible as molecular studies (Yagishita et al. 2002, 2009, Near et al. 2012, 2013) have shown that this would
require a thorough examination of the morphology in Kuhliidae, Microcanthidae, Oplegnathidae, Parascorpididae,
Terapontidae in addition to Girellidae, Kyphosidae and Scorpididae. The morphology of Dichistiidae also shows
affinities with Girellidae, Kyphosidae and Scorpididae (Smith 1935, Leis & van der Lingen 1997), and this family
would also need to be considered for a family designation of Kyphosidae.
The taxonomy of the genus K
yphosus has been confused to the point where many authors did not attempt
species level identification (e.g. Humann & DeLoach 2002). Various genera, subgenera and subspecies have been
proposed for this genus (e.g. Lacepède 1802, 1803, Whitley 1931), and there has been great uncertainty concerning
the taxonomic level of both the family and related genera (e.g. Atypichthys, Girella, Microcanthus and Scorpis)
(Nelson 2006). Currently 16 species of Kyphosidae are recognised (Eschmeyer 2012): Hermosilla azurea,
Kyphosus analogus, K. bigibbus, K. cinerascens, K. cornelii, K. elegans, K. gladius, K. hawaiiensis, K. incisor, K.
lutescens, K. pacificus, K. sectatrix, K. sydneyanus, K. vaigiensis, Neoscorpis lithophilus and Sectator ocyurus.
Delimiting the different species is difficult and is currently based on a combination of number of anal and dorsal
soft fin rays, scale row counts, gill raker counts, colour patterns, relative body proportions, and relative size and
placement of fins. Sakai and Nakabo (1995, 2004, 2006, 2008) considered several species names to be junior
synonyms and limited the number of valid species of Kyphosus to 12 in their revisions of western Indo-Pacific
Kyphosus. Sakai and Nakabo found Pimelepterus lembus, K. bleekeri and K. gibsoni to be junior synonyms of K.
vaigiensis (Sakai & Nakabo 1995), K. fuscus and Pimelepterus fallax to be junior synonyms of K. bigibbus (Sakai
& Nakabo 2004). More recently, Sakai and Nakabo found Pimelepterus indicus, Pimelepterus raynaldi and
Pimelepterus altipinnis to be junior synonyms of K. cinerascens, and pointed out that Segutilum klunzingeri should
be considered a junior synonym of K. sydneyanus (Sakai & Nakabo 2006), and finally that Cantharus lineolatus
was a synonym of K. vaigiensis (Sakai & Nakabo 2008). However, Sakai and Nakabo (1995, 2004, 2006, 2008) did
not consider the possibility that Indo-Pacific Kyphosus might also occur in the Atlantic or the Eastern Pacific, and
their revision of the Indo-Pacific species was based entirely on a morphological examination of valid congeners.
With the recent description of K. gladius
(Knudsen & Clements 2013), the number of valid species of Kypho
sus
immediately predating this revision was 13, a number that was to change following this revision
Kyphosids occur in both the Caribbean Sea and the tropical Central East Pacific Ocean, and geminate kyphosid
species pairs
separated by the Panamanian isthmus (see Craig et al. 2004) were proposed by Jordan (1908) who
regarded the eastern Pacific K. analogus and Atlantic K. incisor to be a species pair. Similarly, Thomson et al.
(2001) regarded the eastern Pacific K. elegans and the Atlantic K. sectatrix as a species pair. No support has yet
been found for these two supposedly geminate species pairs. In this taxonomic revision we looked for
synapomorphic traits among these species pairs to determine if there is any morphological support for these
proposed geminate species pairs.
The morphological differences between N
eoscorpis and Kyphosus can appear quite distinct, e.g. Neoscorpis
has a different arrangement of dorsal- and anal-fin spines and rays (Gilchrist & Thompson 1908, Smith 1931,
Heemstra & Heemstra 2004) to Kyphosus (Sakai & Nakabo 1995, 2006), but both the silvery grey colouration, the
continous dorsal fin, lanceolate teeth and strict herbivory in Kyphosus and Neoscorpis indicate a close relationship
between these two genera, and the common approach is to consider Kyphosus and Neoscorpis members of the same
family (e.g. Heemstra & Heemstra 2004, Nelson 2006, van der Elst & King 2006). In the present revision, we
follow this classification and consider Neoscorpis a genus of Kyphosidae, and thus we have included Neoscorpis in
the identification key. The key to species provided here is largely based on distinctive external characters to
facilitate field identification, but also lists characteristic osteological features.
KNUDSEN & CLEMENTS
6 · Zootaxa 3751 (1) © 2013 Magnolia Press
Materials and methods
Meristic and morphometric characters examined
Morphological measurements and meristics follow Hubbs and Lagler (1958) and Sakai and Nakabo (1995).
Ost
eological characters were examined using more than 450 digital radiographic photos from 35 species
representing Dichistiidae, Girellidae, Kyphosidae, Microcanthidae and Scorpididae. Osteological characters were
recorded following Springer and Smith-Vaniz (2008) and Smith and Bailey (1961). Posterior vertebrae with a
haemal spine were identified as caudal vertebrae. Meristic data were taken on the left side of specimens, however,
for most type material (dependent on the condition of the specimen) both sides were examined. Measurements
were done with a dial calliper to the nearest 0.1 mm and converted to percent of standard length (SL) for
comparison across size ranges. All lengths presented for examined material are given as mm SL. All counts of gill
rakers were performed on the lateral side of the first gill arch, and are presented as counts on the upper and lower
part of the gill arch as well as a total count. Some underwater photographs taken from the right side were reflected
horizontally by using Adobe Photoshop. Material from AIM, AMS, BMNH, BPBM, LACM, MCZ, MNHN,
NZNM, QM, SMNS, WAM, USNM, ZMB and ZMUC was examined for morphological characters. Abbreviations
for museum collections follow Fricke and Eschmeyer (2013).
Geographic distribution presented is based on museum records, DNA sequence from tissue samples and
und
erwater photographic records. The identity of tissue samples was confirmed by molecular markers and by
comparison of morphological characters with type material. The following abbreviations are used for the
morphological traits examined. GRT: Total number of gill rakers on external side of first gill arch facing
operculum. GRU: Number of gill rakers on upper part of gill arch. GRL: Number of gill rakers on lower part of
gill. D: Dorsal fin. A: Anal fin. TScLL: Total number of scales in lateral line, with or without pores. PScLL: Pored
scales in lateral line. ScRLR: Scale rows in longitudinal row. AV: anterior precaudal vertebrae. CV: Caudal
vertebrae. A-ptg: Anal pterygiophores. D-ptg: Dorsal pterygiophores. P1: Pectoral fin. P2: Pelvic fin.
Delineation of morphological groups
Morphological variation was recorded both from voucher specimens used for the molecular analysis (published
separately
) and museum material. The molecular analyses were based on Bayesian analysis (Ronquist et al. 2012)
of both mtDNA and nDNA in a concatenated dataset. Matching vouchered specimens with museum type material
made it possible to identify diagnostic morphological characters for all the species of Kyphosus. The morphological
characters were compared in museum specimens by comparing pairs of species with a one-way ANOVA using the
software PAST (Hammer et al. 2001), and subsequently clusters were inferred from the morphological characters
using a principal component analysis (PCA). These morphological clusters could then be matched to genetic
clusters following the approach for species delimitation described by Feulner et al. (2007). This approach made it
possible to identify key morphological characters that could be included in the identification key.
Taxonomy
Genus Ky
phosus Lacepède
Perca Linnaeus 1758: p. 293. Description of Pimelepterus saltatrix, but in Linnaeus 1766: p. 486 corrected to Perca sectatrix,
based on Catesby (1743) pl. 8, middle figure, no type known. Preoccupied by perches (f. Percidae) (Linnaeus 1758).
Sciæna Forsskål 1775: p. 45. Description of Sciæna cinerascens (valid as Kyphosus cinerascens) from the Red Sea, no type
available. Preoccupied by croakers (f. Sciaenidae) (Linnaeus 1758).
Tahmel Forsskål 1775: p. 45 (originally named Sciena cinerascens, tahmel), considered a synonym of K. cinerascens by Jordan
and Richardson (1908) and an unavailable name by Sakai and Nakabo (2006), no type or diagnosis available.
Kyphosus Lacepède 1801: p. 114, 115 (type locality: Ile de France, Madagascar), holotype: MNHN B-2162 [dried skin]
described as Kyphosus bigibbus by monotypy.
Pimelepterus Lacepède 1802: p. 429 (proposed as subgenus. Monotypic with Pimelepterus bosquii), MNHN 0000-9601 is the
holotype for Pimelepterus bosquii (see Bauchot 1963), although Eschmeyer (2013) incorrectly also listed MNHN 0000-
2977 as a syntype for Pimelepterus bosquii. MNHN 0000-2977 is a holotype of K. sectatrix.
Dorsuarius Lacepède 1803: p. 482 (proposed as subgenus. Monotypic with Dorsuarius nigrescens), the holotype MNHN B-
2162 (dry) is the same specimen as the holotype for K. bigibbus.
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TAXONOMIC REVISION OF KYPHOSIDAE
Xyster Lacepède 1803: p. 484 (proposed as subgenus. Monotypic with Xyster fuscus), the holotype MNHN B-2162 (dry) is the
same specimen as the holotype for K. bigibbus.
Xysterus Rafinesque 1815: p. 88 (incorrect spelling of Xyster).
Seleima Bowdich 1825: p. 238 (proposed as subgenus. Monotypic with Seleima aurata), no type known.
Opisthistius Gill 1862: p. 245 (proposed subgenus for Pimelepterus tahmel and Pimelepterus raynaldi), type by subsequent
designation of Jordan and Evermann 1898 is Sciaena tahmel Forsskål 1775, type as designated by Alleyne and Macleay
1877, holotype: AMS.I 16342-001.
Hermosilla Jenkins and Evermann 1889: p. 144 (proposed as subgenus. Monotypic with Hermosilla azurea), holotype: USNM
39629.
Sectator Jordan and Fesler 1893: p. 533–534, 536 (proposed as subgenus. Monotypic with Sectator ocyurus), holotype:
USNM29395.
Segutilum Whitley 1931: p. 319 (proposed as subgenus. Monotypic with Segutilum klunzingeri) the original designation is
Pimelepterus sydneyanus Günther 1886; Knudsen and Clements (2013) found that the assumed type, SMNS 2673, for
Segutilum cannot be confirmed.
Leptokyphosus Whitley 1931: p. 320 (proposed as subgenus. Monotypic with Kyphosus gibsoni), holotype: QM.I 30.
Cridorsa Whitley 1938: p. 159 (proposed as subgenus. Monotypic with Cridorsa moonta), Glover (1976) noted that the type
SAMA F2023 had been destroyed.
Diagnosis
Mouth terminal. Body oblong, elliptical in lateral view. Teeth lanceolate or incisiform. Dorsal fin with X–XI spines
and 1
0–15 rays. Anal fin with II–III spines and 10–14 rays. Pectoral fin with 15–20 rays. Pelvic fin with I spine and
5 rays. Caudal fin with 16–18 rays. Gill rakers arranged in 3–11, and 11–24 on upper and lower limb, respectively,
giving a total of 15–32 gill rakers. Pored scales in lateral line: 41–64. Total scales in lateral line: 49–82. Scales
along longitudinal row: 44–76. Cheek scales absent to 3–17. Precaudal vertebrae: 10–11, caudal vertebrae: 15–16.
Anal pterygiophores: 11–16. Dorsal pterygiophores: 19–26. Dorsal supraneurals arranged as: 0/0/0–1/… in first to
third dorsal insertion space, occasionally as: 0/0–0/1/... in first to third dorsal insertion space. Dorsal
supranumerary spines associated with first dorsal pterygiophore: 1–2. First dorsal pterygiophore inserts in third
dorsal interhaemal spine space. Anal supranumerary spines associated with first anal pterygiophore: 2. First anal
pterygiophore inserts between the eleventh to twelfth vertebrae. Vertebrae posterior to posteriormost vertebrae
participating in last dorsal and anal interhaemal spine space: 6–9.
Description
Body long, elliptical in lateral view and relatively high, can vary from deep to low (32.7–54.7 %SL), and vary in
wi
dth (10.3–26.4 %SL). Head profile beaked to blunt. Head not comprising more than 38.1 %SL. Six
branchiostegal rays, gill membranes not fused across isthmus. Mouth terminal, horizontal to oblique, with maxilla
partly covered by lachrymal on distal part of maxilla. Upper jaw extending posteriorly to just below anterior edge
of eye or to below centre of eye. Preorbital region without scales. Scales ctenoid and covering the body from the
postorbital region of operculum, and the cheek and posteriorly to the caudal fin. Interorbital region either without
scales or with up to 24 scale rows across. Pelvic fin originates just below pectoral fin or situated well behind
anterior part of pectoral fin (1.9–16.5 %SL). Caudal fin deeply emarginated to truncate.
Colour when fresh—v
ariable across the genus. Grey to silvery, bronze or olive green on a silvery background,
or with 5–8 broad, vertical dark bands. Some species have horizontal golden lines of scales along the body from
operculum to caudal fin, or with broad shiny blue and yellow lines horizontally along the body from operculum to
caudal fin. One species has a completely yellow (xanthic) variant. Juveniles often mottled or with vague spots. A
spotted pattern is also sometimes displayed in adults, and is more obvious when associated with aggressive
behaviour. Some species have a pronounced white or silver horizontal streak below eye.
Genus Neoscorpis Smith
Scorpis Gilchrist and Thompson 1908: p. 162 (Natal coast South Africa), described as Scorpis lithophilus, holotype: SAM
9926; Gilchrist and Thompson 1917: p. 365.
Neoscorpis Smith 1931: p. 8 (monotypic with N. lithophilus), holotype: SAM 9926; Smith 1961: p. 249, pl. 34; van der Elst
1981: p. 282; Smith 1986: p. 604, pl. 68 (listed under Scorpididae); Pereira 2000: p. 11 (Mozambique, listed under
Scorpididae); Heemstra and Heemstra 2004: p. 251; van der Elst and King 2006: p. 73.
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8 · Zootaxa 3751 (1) © 2013 Magnolia Press
Diagnosis
Mouth terminal. Body oblong, elliptical in lateral view. Teeth lanceolate or incisiform. Dorsal fin with VI–VIII
spines a
nd 20–23 rays, anal fin with III spines and 23–26 rays. Pectoral fin with 16–18 rays. Pelvic fin with I spine
and 5 rays. Gill rakers arranged in 6–8, and 12–15 on upper and lower limb, respectively, giving a total of 20–23
gill rakers. Pored scales in lateral line: 41–64. Total scales in lateral line: 90–97. Scales along longitudinal row: 72–
76. Interorbital scales absent (Smith 1931, 1986, Heemstra & Heemstra 2004).
Description
Body long, elliptical in lateral view and relatively high. Young specimens quite compressed, older individuals
rel
atively broad. Snout as long as eye and quite blunt. Mouth terminal, horizontal to oblique. Anterior soft rays in
dorsal and anal fins markedly extended. Upper jaw extending posteriorly to just below anterior edge of eye or to
below centre of eye. Scales ctenoid covering the body from the postorbital region of operculum and the cheek
posteriorly to the caudal fin. Pelvic fin originates well behind anterior part of pectoral fin. Caudal fin emarginate
(Gilchrist & Thompson 1908, Smith 1931, Heemstra & Heemstra 2004).
Colour when fresh—Grey
to silvery body, fins darker and more blue in colour. Blueish colour on anterior-
dorsal part of body. Edge of operculum dark. Juveniles with up to eight vertical faint dark bars.
Remarks
Neoscorpis lithophilus was orig
inally considered a species within Scorpis (Gilchrist & Thompson 1908). Smith
(1931), however, found that lower number of dorsal fin spines, the absence of a serrated edge on the preoperculum,
and the difference in teeth-shape did not match Scorpis, and therefore erected Neoscorpis, monotypic with N.
lithophilus. Later, Nelson (2006) assigned Neoscorpis to Kyphosinae. A close relationship between Neoscorpis and
Kyphosus was corroborated by our molecular analysis (published separately), suggesting that the similarities
between Neoscorpis and Scorpididae are plesiomorphic.
Key to the species of Kyphosidae:
1. Fewer than IX dorsal fin spines (VI–VIII); more than 19 soft dorsal-fin rays (20–23); more than 22 soft anal-fin rays (23–26),
anterior part of soft dorsal- and soft anal fin distinctly elongate (South Africa, southern Mozambique) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoscorpis lithophilus Stone bream
More than IX dorsal fin spines (X–XI); fewer than 17 soft dorsal-fin rays (10–16); fewer than 16 soft anal-fin rays (10–15),
anterior part of soft dorsal- and soft anal fin not distinctly elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Scales absent on interorbital region; fewer than 9 postorbital scales (5–8); 25 vertebrae in total . . . . . . . . . . . . . . . . . . . . . . . . 3
Scales present on interorbital region; more than 9 postorbital scales (10–21); 26 vertebrae in total . . . . . . . . . . . . . . . . . . . . . . 4
3.
Caudal fin with white edges; 14–16 soft dorsal fin rays; more than 12 soft anal fin rays (13–15); no distinct vertical bands
across body (Western Australia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus cornelii Buffalo bream
Caudal fin without white edges; 10–11 soft dorsal fin rays; less than 11 soft anal fin rays (10); 5–6 vertical dark green bands
across body (southern California to Baja). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus azureus Zebraperch
4. More than 21 (occasionally 19) total gill rakers (19–32); more than 56 total scale rows along the lateral line (usually more than
65) (56–85); often more than 70 scale rows along longitudinal row (50–77); unpaired fins not dusky . . . . . . . . . . . . . . . . . . . . 5
Fewer than 21 (occasionally with up to 22) total gill rakers (15–22); fewer than 76 total scale rows along lateral line (usually
less than 66) (52–76); fewer than 68 scale rows along longitudinal row (48–67); unpaired fins dusky . . . . . . . . . . . . . . . . . . 10
5.
No more than 12 soft dorsal fin rays (occasionally 13) (11–13); 10–13 soft anal-fin rays (usually 11) (10–13); never more than
33 pterygiophores in total (32–33); not more than 70 scale rows along longitudinal row (50–69); soft rayed part of dorsal and
anal fins often highly elevated in individuals with 13 dorsal fin rays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Thirteen or more soft dorsal fin rays (13–15); 12–14 soft anal fin rays; 34 or more pterygiophores in total (34–39); often more
than 70 scale rows along longitudinal row (52–77); soft rayed part of dorsal and anal fin not elevated . . . . . . . . . . . . . . . . . . . 8
6. Soft-rayed part of dorsal and anal fins extended; fourth dorsal fin ray long (8.7–19.1 %SL); second anal fin ray long (10.4–22.3
%SL); 25 or more total gill rakers (25–30) (Atlantic Ocean, Indo-Pacific Ocean) . . . . . . Kyphosus cinerascens Highfin chub
Soft-rayed part of dorsal- and anal-fin not extended; fourth dorsal fin ray short (4.8–12.0 %SL); second anal fin ray short (8.5–
16.1 %SL); 25 or fewer total gill rakers (19–25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. More than 21 total gill rakers (22–25); pelvic fin short (11.8–19.0 %SL); second anal fin ray short (8.5–15.2 %SL) (Atlantic
Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus sectatrix Beaked chub
Usually fewer than 23 (never more than 22) total gill rakers (19–22); pelvic fin long (19.3–20.8 %SL); second anal fin ray long
(13.5–16.1 %SL) (Hawaiian Islands) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus hawaiiensis Hawaiian chub
8.
Soft part of dorsal fin with 13 rays; anal fin with 12 soft rays; interorbital width 11.0–13.8 %SL; 21–22 dorsal pterygiophores;
no more than 13 anal pterygiophores; 52–57 pored scales in lateral line; not more than 69 total scales along lateral line (61–
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TAXONOMIC REVISION OF KYPHOSIDAE
69); relative low number scale rows in the longitudinal row (52–62); body colour uniform grey or bronzed brown or mottled
with white spots; no conspicuous golden horizontal scale rows along body (east Pacific) . . . . Kyphosus elegans Cortez chub
Soft part of dorsal fin with 13 or more rays (13–15); anal fin with 12–15 soft rays; interorbital width 9.6–14.2 %SL; more than
22 dorsal pterygiophores (22–24); 13 or more anal pterygiophores (13–15); 52–64 pored scales in lateral line; regularly with
70 or more scales in total along lateral line (63–85); relative high number scale rows in the longitudinal row (56–77); body
with either clear golden horizontal scale rows or with large distinct yellow and blue wavy stripe horizontally along body. . . . 9
9.
Soft part of dorsal fin with 13–14 rays; soft part of anal fin with 11–14 rays; 22–24 dorsal pterygiophores; not more than 14
anal pterygiophores (13–14); golden yellow horizontal scale rows along body from operculum to caudal fin; caudal fin not
deep emarginated (Atlantic Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus vaigiensis Lowfin chub
Soft part of dorsal fin with 15 rays or more (15–16); soft part of anal fin with 14 rays or more (14–15); 24 dorsal pterygio-
phores; 15 anal pterygiophores; a blue and yellow wavy stripe horizontally across body from operculum to caudal fin; blue
s
treak behind eye; caudal fin deeply emarginated (Pacific Ocean). . . . . . . . . . . . . . . . . . . . . Kyphosus ocyurus Rainbow chub
10. Pored scales in lateral line 51–61; scales in longitudinal row 54–67; total gill rakers 18–22; anal fin base long (18.1–24.8
%SL); line along ventral edge of anal fin extended backwards through caudal peduncle follows dorsal edge of caudal fin
(Atlantic Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus bigibbus Darkfin chub
Pored scales in lateral line 42–56; scales in longitudinal row 48–57; total gill rakers 15–20; anal fin base short (14.0–22.4 %SL);
line along ventral edge of anal fin extended backwards through caudal peduncle cuts above dorsal edge of caudal fin. . . . . . . 11
11. Vertebrae 10 precaudal +16 caudal; caudal peduncle not deep (9.9–11.8 %SL); sixth dorsal-fin ray length short (8.7–11.8
%SL); pelvic fin short (13.4–16.2 %SL); green vertical bar on operculum; line along ventral edge of anal fin runs parallel to
the dorsal edge of the caudal fin (west Australia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus gladius Gladius drummer
Vertebrae 11 precaudal +15 caudal; caudal peduncle deep (12.1–14.5 %SL); sixth dorsal-fin ray length long (11.4–16.1 %SL);
pelvic fin long (14.7–22.5 %SL); no green bar on operculum; line along ventral edge of anal fin does not run parallel to the
dorsal edge of the caudal fin (southern Australia, northern New Zealand) . . . . . . . . . .Kyphosus sydneyanus Silver drummer
Regional keys to the species of Kyphosidae:
To simplify identification we include separate regional keys for the Atlantic Ocean, the Indo-Pacific, and the east
Pacific. However, we stress the importance of using these regional keys with caution, as many species of Kyphosus
have wide and poorly documented distributions, and past misidentification has often resulted from underestimating
the range of some species. The use of these regional keys assumes that our knowledge of species distribution is
complete, and this is unlikely.
Key to the Atlantic species of Kyphosidae:
1. More than 22 (occasionally 22) total gill rakers (22–32); more than 56 total scale rows along the lateral line (usually more than
65) (56–85); often more than 70 scale rows along longitudinal row (50–77); unpaired fins not dusky . . . . . . . . . . . . . . . . . . . . 2
Fewer than 22 (occasionally with up to 22) total gill rakers (18–22); fewer than 76 total scale rows along lateral line (usually
less than 66) (61–76); fewer than 68 scale rows along longitudinal row (54–67); unpaired fins dusky (Atlantic Ocean, Indo-
Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus bigibbus Darkfin chub
2.
No more than 12 soft dorsal fin rays (occasionally 13) (11–13); 10–13 soft anal-fin rays (usually 11) (10–13); never more than 33
pterygiophores in total (32–33); not more than 70 scale rows along longitudinal row (50–69); soft rayed part of dorsal and anal
fins often highly elevated in individuals with 13 dorsal fin rays; no conspicuous golden horizontal scale rows along body . . . . 3
Thirteen or more soft dorsal fin rays (13–15); 11–14 soft anal fin rays; 36 or more pterygiophores in total (36–38); often more than
70 total scales rows along lateral line (63–80); soft rayed part of dorsal and anal fin not elevated; golden yellow horizontal scale
rows along body from operculum to caudal fin (Atlantic Ocean, Indo-Pacific Ocean) . . . . . . . Kyphosus vaigiensis Lowfin chub
3.
Soft-rayed part of dorsal and anal fins extended; fourth dorsal fin ray long (8.7–19.1 %SL); second anal fin ray long (10.4–22.3
%SL); 25–30 total gill rakers (Atlantic Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . . Kyphosus cinerascens Highfin chub
Soft-rayed part of dorsal- and anal-fin not extended; fourth dorsal fin ray short (4.8–11.9 %SL); second anal fin ray short (8.5–
15.2 %SL); 22–25 total gill rakers (Atlantic Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . Kyphosus sectatrix Beaked chub
Key to the Indo-Pacific species of Kyphosidae:
1. Fewer than IX dorsal fin spines (VI–VIII); more than 19 soft dorsal-fin rays (20–23); more than 22 soft anal-fin rays (23–26)
(South Africa, southern Mozambique), anterior part of soft dorsal- and soft anal fin distinctly elongate . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoscorpis lithophilus Stone bream
More than IX dorsal fin spines (X–XI); fewer than 17 soft dorsal-fin rays (10–16); fewer than 16 soft anal-fin rays (10–15),
anterior part of soft dorsal- and soft anal fin not distinctly elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
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10 · Zootaxa 3751 (1) © 2013 Magnolia Press
2. Scales absent on interorbital region; fewer than 9 postorbital scales (5–8); 25 vertebrae in total; caudal fin with white upper and
lower edges; 14–16 soft dorsal fin rays (Western Australia) . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus cornelii Buffalo bream
Scales present on interorbital region; more than 9 postorbital scales (10–21); 26 vertebrae in total; caudal fin without white
edges; not more than 15 soft dorsal fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3.
More than 21 (occasionally only 19) total gill rakers (19–32); more than 56 total scale rows along the lateral line (usually more
than 65) (56–85); often more than 70 scale rows along longitudinal row (50–76); unpaired fins not dusky . . . . . . . . . . . . . . . . 4
Fewer than 21 (occasionally with up to 22) total gill rakers (15–22); fewer than 76 total scale rows along lateral line (usually
less than 65) (52–76); fewer than 70 scale rows along longitudinal row (49–70); unpaired fins dusky . . . . . . . . . . . . . . . . . . . 8
4. No more than 12 soft dorsal fin rays (occasionally 13) (11–13); 10–13 soft anal-fin rays (usually 11) (10–13); never more than
33 pterygiophores in total (32–33); not more than 68 scale rows along longitudinal row (45–68); often less than 70 scales in
total along lateral line (56–81); soft rayed part of dorsal and anal fins often highly elevated in individuals with 13 dorsal fin
rays; no conspicuous golden horizontal scale rows along body and no distinct yellow and blue wavy stripe on body . . . . . . . . 5
Thirteen or more soft dorsal fin rays (13–15); 12–14 soft anal fin rays; 36 or more pterygiophores in total (36–39); often more
than 70 scale rows along longitudinal row (59–77); regularly with 70 or more scales in total along lateral line (63–85); soft
rayed part of dorsal and anal fin not elevated; body with either clear golden horizontal scale rows or with large distinct yellow
and blue wavy stripe horizontally along body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5.
Soft-rayed part of dorsal and anal fins extended; fourth dorsal fin ray long (8.7–19.1 %SL); second anal fin ray long (10.4–22.3
%SL); 25 or more total gill rakers (25–30) (Atlantic Ocean, Indo-Pacific Ocean) . . . . . . Kyphosus cinerascens Highfin chub
Soft-rayed part of dorsal- and anal-fin not extended; fourth dorsal fin ray short (4.8–12.0 %SL); second anal fin ray short (8.5–
16.1 %SL); 25 or fewer total gill rakers (22–25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6.
More than 21 total gill rakers (22–25); pelvic fin short (11.8–19.0 %SL); second anal fin ray short (8.5–15.2 %SL) (Atlantic
Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus sectatrix Beaked chub
Fewer than 23 total gill rakers (19–22); pelvic fin long (19.3–20.8 %SL); second anal fin ray long (13.5–16.1 %SL) (Hawaiian
Islands) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus hawaiiensis Hawaiian chub
7.
Soft part of dorsal fin with 13–14 rays; soft part of anal fin with 11–14 rays; not more than 21 dorsal pterygiophores (22–24);
not more than 14 anal pterygiophores (12–14); golden yellow horizontal scale rows along body from operculum to caudal fin;
caudal fin not deep emarginated (Atlantic Ocean, Indo-Pacific Ocean). . . . . . . . . . . . . . . . . Kyphosus vaigiensis Lowfin chub
Soft part of dorsal fin with 15 rays; soft part of anal fin with 14 rays; 24 dorsal pterygiophores; 15 anal pterygiophores; a blue
and yellow wavy stripe horizontally across body from operculum to caudal fin; blue streak behind eye; caudal fin deeply emar-
ginated (Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus ocyurus Rainbow chub
8.
Pored scales in lateral line 51–61; scales in longitudinal row 54–67; total gill rakers 18–22; anal fin base long (18.1–24.8
%SL); line along ventral edge of anal fin extended backwards through caudal peduncle follows dorsal edge of caudal fin
(Atlantic Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus bigibbus Darkfin chub
Pored scales in lateral line 42–56; scales in longitudinal row 48–57; total gill rakers 15–20; anal fin base short (14.0–22.4 %SL);
line along ventral edge of anal fin extended backwards through caudal peduncle cuts above dorsal edge of caudal fin. . . . . . . . 9
9. Vertebrae 10 precaudal +16 caudal; caudal peduncle not deep (9.9–11.8 %SL); sixth dorsal-fin ray length short (8.7–11.8
%SL); pelvic fin short (13.4–16.2 %SL); green vertical bar on operculum; line along ventral edge of anal fin runs parallel to
the dorsal edge of the caudal fin (west Australia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus gladius Gladius drummer
Vertebrae 11 precaudal +15 caudal; caudal peduncle deep (12.1–14.5 %SL); sixth dorsal-fin ray length long (11.4–16.1 %SL);
pelvic fin long (14.7–22.5 %SL); no green bar on operculum; line along ventral edge of anal fin does not run parallel to the
dorsal edge of the caudal fin (southern Australia, northern New Zealand) . . . . . . . . . .Kyphosus sydneyanus Silver drummer
Key to the East Pacific species of Kyphosidae:
1. Scales absent on interorbital region; fewer than 9 postorbital scales (7–8); 25 vertebrae in total; 10–11 soft dorsal fin rays; 5–6
vertical dark green bands across body (southern California to Baja) . . . . . . . . . . . . . . . . . . . . . Kyphosus azureus Zebraperch
Scales present on interorbital region; more than 9 postorbital scales (10–21); 26 vertebrae in total; 11 or more soft dorsal fin
rays (11–15); no distinct vertical green bands across body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. No more than 12 soft dorsal fin rays (11–12); 10–12 soft anal-fin rays (usually 11); never more than 33 pterygiophores in total
(32–33); not more than 69 scale rows along longitudinal row (60–69); more than 21 total gill rakers (22–25); body colour can
be grey without obvious golden horizontal scale rows or be wholly or partly yellow (xanthic) (Atlantic Ocean, Indo-Pacific
Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus sectatrix Beaked chub
Thirteen or more soft dorsal fin rays (13–15); 12–14 soft anal fin rays; 34 or more pterygiophores in total (34–39); often more
than 70 scale rows along longitudinal row (52–77); 23–32 total gill rakers; body colour can be bronzed or brown and can have
white blotches, or the body is grey with golden horizontal scale rows, or partly yellow and blue and have a yellow wavy stripe
horizontally across body from operculum to caudal fin and blue streak behind eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Soft part of dorsal fin with 13 rays; anal fin with 12 soft rays; interorbital width 11.0–13.8 %SL; 21–22 dorsal pterygiophores;
13 anal pterygiophores; 52–57 pored scales in lateral line; not more than 69 total scales along lateral line (61–69); relative low
number scale rows in the longitudinal row (52–62); body colour uniform grey or bronzed brown or mottled with white spots;
no conspicuous golden horizontal scale rows along body (east Pacific) . . . . . . . . . . . . . . . . . . Kyphosus elegans Cortez chub
Soft part of dorsal fin with 13 or more rays (13–15); anal fin with 12–15 soft rays; interorbital width 9.6–14.2 %SL; more than
22 dorsal pterygiophores (22–24); 13 or more anal pterygiophores (13–15); 52–64 pored scales in lateral line; regularly with
Zootaxa 3751 (1) © 2013 Magnolia Press · 11
TAXONOMIC REVISION OF KYPHOSIDAE
70 or more scales in total along lateral line (63–85); relative high number scale rows in the longitudinal row (56–77); body
with either clear golden horizontal scale rows or with large distinct yellow and blue wavy stripe horizontally along body. . . . 4
4. Soft part of dorsal fin with 13–14 rays; soft part of anal fin with 11–14 rays; 22–24 dorsal pterygiophores; 12–14 anal pterygio-
phores; golden yellow horizontal scale rows along body from operculum to caudal fin; caudal fin not deep emarginated (Atlan-
tic Ocean, Indo-Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyp
hosus vaigiensis Lowfin chub
Soft part of dorsal fin with 15 rays or more; soft part of anal fin with 14 rays or more; 24 dorsal pterygiophores; 15 or more
anal pterygiophores; a blue and yellow wavy stripe horizontally across body from operculum to caudal fin; blue streak behind
eye; caudal fin deeply emarginated (Pacific Ocean) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus ocyurus Rainbow chub
Kyphosus azureus (Jenkins & Evermann, 1889)
(Common name: zebraperch, zebra-striped sea perch, chopa bonita)
(Figure 1, 3, 4 Table 1, 8–18)
Hermosilla azurea Jenkins and Evermann 1889: p. 144–145 (type-locality: Guaymas, Sonora, western Mexico), holotype:
USNM 39629; Thomson et al. 1979: p. 123 (Monterey, California, Sea of Cortez); Eschmeyer and Herald 1983: p. 224, pl.
34; Goodson 1988: p. 89; Stevens et al. 1989: p. 759, fig. 1, 6, 7; Robins et al. 1991: p. 55 (Pacific); Allen and Robertson
1994: p. 176 (Baja California, Californian Gulf); Sommer 1995: p. 1199 (key) (Californian Gulf); De La Cruz Agüero et
al. 1997: p. 194 (Baja California, Californian Gulf); Gotshall 1998: p. 47, fig. 101 (California); Thomson et al. 2001: p.
141; Sturm and Horn 2001: p. 170 (Southern Californian Bight); Nelson et al. 2004: p. 150; Fidopiastis et al. 2006: p. 631.
Hermosilla robusta Osburn and Nichols 1916: p. 166, fig. 11 (type-locality: Tiburon Island, Gulf of California, Mexico),
holotype: USNM 87547; Thomson et al. 2001: p. 141.
Material examined
5 specimens (33–306 mm SL): USNM 321281, (2 specimens: 33–45 mm), Mexico, Isla Sal St. Pudes; USNM
288
876, 69 mm, Mexico, Baja California, Baja Isla Cedros; USNM 39629 (holotype of H. azurea), 155 mm,
Guaymas, Sonora, Mexico; USNM 87547 (holotype of H. robusta), 306 mm, Tiburon Island, Gulf of California,
Mexico.
FIGURE 1. Museum material and freshly caught specimens of Kyphosus azureus and K. cornelii, photos by SWK unless
specified otherwise, all lengths are in mm SL, scale bars equal 10 mm: A) Kyphosus azureus, USNM 39629, 155 mm, holotype
of Hermosilla azurea (photo by Sandra J. Raredon, Smithsonian Institution), B) Kyphosus azureus, USNM 87547, 306 mm,
holotype of Hermosilla robusta (photo by Sandra J. Raredon, Smithsonian Institution), C) Kyphosus cornelii, WAM P. 25853-
009, 196 mm, D) Kyphosus cornelii, WAM P.33451-003, 174 mm, neotype of Kyphosus cornelii.
Diagnosis
Interorbital region without scales. Mouth terminal. Teeth incisiform. Body with 5–9 vertical dark bands. Dorsal fin
wi
th XI spines and 10–11 rays, anal fin with III spines and 10 rays. Gill rakers arranged 4–5, and 11–12 on upper
KNUDSEN & CLEMENTS
12 · Zootaxa 3751 (1) © 2013 Magnolia Press
and lower limb, respectively. Lateral line with 49–55 scales in total, of which 45–49 have pores. Longitudinal row
with 43–49 scales. Cheek scales 5–6. Postorbital scales 7–8. Precaudal vertebrae 10, caudal vertebrae 15. Dorsal
and anal pterygiophores 19–20 and 11, respectively.
FIGURE 2. Museum material and freshly caught specimens of Kyphosus bigibbus, photos by SWK unless specified otherwise,
all lengths are in mm SL, scale bars equal 10 mm: A) SMNS 23084-1, 102 mm, paralectotype of Pimelepterus fallax, B) SMNS
23084-2, 91 mm, paralectotype of Pimelepterus fallax, C) MNHN 0000-9601, 123 mm, holotype of Pimelepterus bosquii, D)
MNHN B-2162, >303 mm, holotype of Kyphosus bigibbus, E) NMNZ P.34930, 176 mm (photo by Carl Struthers, NMNZ), F)
WAM P.33451-001, 270 mm, G) AIM MA30189, 219 mm, H) ZMB 7997, 186 mm.
Description
Type material.
Holotype (USNM 39629): D: XI, 11; A: III,
10; P1: 16/16; P2: I, 5; PScLL: 49/47; TScLL: 55/53; ScRLR: 47/45;
GRU: 4/4; GRL: 11/12; GRT: 15/16; PCV: 10; CV: 15; D-Ptg: 20, A-Ptg: 11.
Holotype of Hermosilla robusta (USNM 87547): D: XI, 10; A: III, 10; P1: 16; P2: I, 5; PScLL: 49; TScLL: 51;
ScRLR: 48; GRU: –/–; GRL: –/–; GRT: –/–; PCV: 10; CV: 15; D-Ptg: 19, A-Ptg: 11.
Other material (including type material): D: XI, 10–11; A: III, 10; P1: 15–17; P2: I, 5; PScLL: 45–49; TScLL: 49
55; ScRLR: 43–49; GRU: 4–5; GRL: 11–12; GRT: 15–17; PCV: 10; CV: 15; D-Ptg: 19, A-Ptg: 11.
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TAXONOMIC REVISION OF KYPHOSIDAE
FIGURE 3. Underwater photographs of three species of Kyphosus, photos by KDC unless specified otherwise: A) Kyphosus
azureus, California (photo by Daniel Gotshall), B) Kyphosus bigibbus, Poor Knights Island, New Zealand, December 2009, C)
Kyphosus bigibbus, Rottnest Island, Western Australia, July 2011, D) Kyphosus bigibbus, Rottnest Island, Western Australia,
July 2011, E) Kyphosus cornelii, Rottnest Island, Western Australia, July 2011, F) Kyphosus cornelii, Rottnest Island, Western
Australia, July 2011, G) Kyphosus bigibbus, near Poor Knights Island, New Zealand, December 2009, H) Kyphosus bigibbus,
Rottnest Island, Western Australia, May 2011.
Body long, elliptical in lateral view. Caudal fin not deeply emarginated. Body relatively deep (36.2–50.6
%SL), but not wide (13.7–16.2 %SL). Head short (26.6–33.0 %SL), in profile, sloping gently from above eye,
giving the snout a pointed appearance. Snout (6.0–8.1 %SL) equals eye diameter in length (7.0–11.0 %SL).
Interorbital region narrow (10.9–11.9 %SL). Caudal peduncle relatively deep (13.1–14.9 %SL) and long (20.4–
24.1 %SL). Scales ctenoid, fairly large, and present behind preopercular region and covering the posterior part of
the body to the caudal fin. Scales absent on interorbital region. Mouth terminal and ventrally oblique. Head profile
slightly pointed, slanting abruptly from interorbital region to snout. Teeth incisor-shaped arranged in a single row
on anterior edge of dentary and premaxilla, increasing in numbers in adult individuals. Smaller patch of tiny cone
shaped teeth arranged in 3–4 rows in mouth floor and on mouth roof. Anal fin base relatively short (14.0–18.9
%SL), compared to the length of the dorsal fin base. Spinous part of dorsal fin higher (sixth dorsal fin spine 13.8–
15.1 %SL) than soft-rayed part of dorsal fin (fourth dorsal fin ray 8.6–13.0 %SL). Spinous part of dorsal fin longer
KNUDSEN & CLEMENTS
14 · Zootaxa 3751 (1) © 2013 Magnolia Press
(19.9–31.4 %SL) than soft-rayed part of dorsal fin (15.7–21.1 %SL). Posterior edges of soft-rayed part of anal- and
dorsal fin rounded. Anal fin low, with second soft ray attaining longest length (10.2–16.6 %SL) of anal fin rays.
Pectoral fin with rounded posterior edge and as long (19.3–21.9 %SL) as pelvic fin (15.2–19.6 %SL). Anal fin
placed moderately further back, giving a relatively long pre-anal length (57.6–67.3 %SL).
Colour when fresh
greenish to silvery, with five to nine broad olive-green vertical bars. Head uniform brown
to green. Ventral body pale or white and silvery. Dorsal and caudal fins dark, and slightly darker at the edges of the
fins towards the rear of the fin. Base of pectoral- and pelvic fin white or silvery. Eye with yellow iris and black
pupil. Occasionally with a dark spot at posterior-ventral corner of pectoral fin base (Figure 1), and with dark spot
midlaterally on posterior edge of preoperculum.
Colour of preserved specimens—silv
ery to grey with five to seven brown or dark vertical bars, or entire body
uniform brown (Figure 1 and 3). The black mark at the ventral edge of the pectoral fin base and the black mark
midlaterally on the posterior edge of the preoperculum is not necessarily consistent as it can fade in preserved
specimens.
Distribution
Found in temperate to subtropical seas, from Monterey on the Pacific coast of California and south to Baja
C
alifornia and in the Sea of Cortez (Figure 4). More common in the northern and central part of the Sea of Cortez
(Thomson et al. 2001), but this could be changing with an increase in numbers recorded 10 years ago in the
Southern California Bight (Sturm & Horn 2001). Inhabits coastal rocky reefs and reef flats with algal growth, in
shallow waters from the surface and down to 10 m, both as solitary individuals and schooling with other species
(De La Cruz Agüero et al. 1997, Sturm & Horn 2001). Exclusively herbivorous (Fidopiastis et al. 2006).
Remarks
Jenkins and Evermann (1889) described Hermosil
la azurea and noted the similarities with Kyphosus, but separated
Hermosilla from Kyphosus on the basis of weaker gill-rakers, margin of preoperculum entire, teeth absent from
tongue and vomer, no band of villiform teeth behind incisor teeth, larger scales, around 55 scales in longitudinal
series, and head, snout, preorbitals, chin, and space below eye without scales. Jenkins and Evermann (1889) found
that Hermosilla differed from Kyphosus in the relative proportions of the spinous and soft portions of the dorsal fin
(spinous portion longer than soft portion), and in the soft part of the anal fin, which is shorter than the soft part of
the dorsal fin. Monotypic with H. azurea, however, our molecular data (published separately) indicates a sister-
species relationship with K. cornelii. A comparison of material of K. cornelii from Western Australia with that of
Hermosilla from California indicates that both H. azurea and K. cornelii lack scales in the interorbital region and
on the preoperculum (where other species of Kyphosus have scales dorsally on the interorbital region and the
preoperculum), and together with K. gladius and K. sydneyanus, both H. azurea and K. cornelii have a relatively
low number of total gill rakers (i.e. H. azurea and K. cornelii have 15–21 gill rakers in total, and K. gladius and K.
sydneyanus have 15–20 gill rakers in total) in comparison with other species of Kyphosus (19–32). Hermosilla
azurea and K. cornelii also share a low number of vertebrae (25 vertebrae in total in both H. azurea and K. cornelii)
in comparison with other species of Kyphosus (26 vertebrae in total) (Table 3). Hermosilla azurea and K. cornelii
both have large scales, which make the total number of scales in longitudinal row (50–59) low when compared to
other species of Kyphosus (up to 76 scales in longitudinal row) (Table 3). We therefore consider the genus
Hermosilla a junior synonym of Kyphosus, and accordingly the species becomes Kyphosus azureus for consistency
of gender. Little is known about the larvae and juveniles, but pelagic juveniles have been reported to be mottled and
with a pattern of pale spots (Stevens et al. 1989), which agrees with other species of Kyphosus that have juveniles
exhibiting a similar spotted phase. Hermosilla robusta was described from Port San, Lower California by Osburn
and Nichols (1916), and diagnosed as differing from H. azurea by having a deeper body, stouter gill rakers and
ventral fins not extending as far back as in H. azurea. Examination of the type specimen of H. robusta (USNM
87547) did not indicate that these proportions vary from the other specimens of H. azurea. No differentiation in
population structure is evident from the mitochondrial DNA control region (Bernardi
et al. 20
03), supporting
dispersal abilities previously recorded for Kyphosus and Hermosilla (Stevens et al. 1989, Sturm & Horn 2001). The
zebraperch is strictly herbivorous, and displays hindgut fermentation (Fidopiastis et al. 2006) much like the
hindgut fermentation found in other Kyphosus (Rimmer 1986, Rimmer & Wiebe 1987, Clements & Choat 1997,
Moran & Clements 2002, Mountfort et al. 2002).
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TAXONOMIC REVISION OF KYPHOSIDAE
FIGURE 4. Map showing locations for samples and museum material of K. azureus (triangles), K. hawaiiensis (diamonds) and K. cornelii (circles) examined in this study. Kyphosus
azureus museum specimens (black triangles), K. azureus only tissue samples without specimens (white triangles), K. azureus type-specimens (red triangles), K. cornelii museum
specimens (black circles), K. cornelii only tissue samples without specimens (white circles), K. cornelii tissue samples with specimens (yellow circles), K. cornelii type-specimens
(red circles), K. hawaiiensis museum specimens (black diamonds), K. hawaiiensis only tissue samples without specimens (white diamonds), K. hawaiiensis tissue samples with
specimens (yellow diamonds), K. hawaiiensis type-specimens (red diamonds).
KNUDSEN & CLEMENTS
16 · Zootaxa 3751 (1) © 2013 Magnolia Press
TABLE 1.
Counts and measurements of
Kyp
hosus azureus and
Kyp
hosus bi
g
ibbus. Values in
p
arentheses indicate mean and standard deviation.
Kyphosus azureus Kyphosus bigibbus
USNM 39629
H
ermosilla azurea
(HT)
USNM 87547
H
ermosilla robusta
(HT)
Range of character, for
specimens examined,
including holotype, paratypes
and non-types.
n
MNHN B–2162
K
yphosus bigibbus
(HT)
SMNS 23084–1
P
imelepterus fallax
(PLT)
SMNS 23084–2
P
imelepterus fallax
(PLT)
MNHN 0000–9601
P
imelepterus bosquii
(HT)
Range of character, for
specimens examined,
including holotype, paratypes
and non-types.
n
Total length (mm) 185 353 39–353 (142.7 ± 130.7) 5 >350 121 107 145 70–595 (260.8 ± 125.2) 56
Standard length (mm) 155 306 33–306 (121.7 ± 113.3) 5 >298 102 91 123 62–527 (219.6 ± 109.8) 56
Counts
Dorsal fin spines XI XI XI (11.0 ± 0.0) 5 XI XI XI XI XI (11.0 ± 0.0) 56
Soft dorsal fin rays 11 10 10–11 (10.8 ± 0.4) 5 12 12 12 12 11–12 (12.0 ± 0.2) 54
Anal fin spines III III III (3.0 ± 0.0) 5 III III III III III (3.0 ± 0.0) 55
Soft anal fin rays 10 10 10 (10.0 ± 0.0) 5 11 11 11 11 10–12 (11.1 ± 0.4) 53
Pectoral fin rays 16/16 16/– 16 (16.0 ± 0.0) 5 –/18 19/19 19/19 –/19 16–20 (18.2 ± 0.9) 52
Pelvic fin rays I, 5 I, 5 (5.0 ± 0.0) 5 I, 5 I, 5 I, 5 I, 5 (5.0 ± 0.0) 54
Caudal fin rays 17 17 17 (17.0 ± 0.0) 5 17 17 17 17 17–18 (17.0 ± 0.2) 48
Gill rakers upper limb on first gill arch 4/4 4–5 (4.3 ± 0.5) 4 6/5 4/4 6/6 4–7 (5.1 ± 0.8) 53
Gill rakers lower limb on first gill arch 11/12 11–12 (11.5 ± 0.6) 4 14/14 14/14 15/15 13–17 (14.2 ± 0.9) 52
Gill rakers, total on first gill arch 15/16 15–17 (15.8 ± 1.0) 4 20/19 18/18 21/21 18–22 (19.3 ± 1.2) 52
Pored scales in lateral line 49/47 49/– 45–49 (47.6 ± 1.7) 5 –/56 59/60 56/54 58/58 51–60 (55.6 ± 2.3) 53
Pored scales in caudal fin 7/– 4/– 4–7 (5.0 ± 1.2) 5 –/– 5/ 3/– 5/– 2–8 (5.6 ± 1.4) 45
Scale rows above lateral line 10/– 10/– 10 (10.0 ± 0.0) 5 –/12 14/13 12/– 12/– 10–15 (12.3 ± 1.1) 54
Scale rows below lateral line 15/16 17/– 15–17 (15.6 ± 0.9) 5 –/22 19/20 19/20 19/21 16–24 (20.8 ± 1.6) 55
Total scales in lateral line 55/53 51/– 49–55 (51.6 ± 2.4) 5 –/74 65/67 67/66 72/74 61–76 (68.9 ± 3.8) 55
Scale rows in longitudinal row 47/45 48/– 43–49 (47.0 ± 1.9) 5 –/66 57/59 64/63 66/65 54–67 (61.7 ± 3.2) 55
Postorbital scales 7/7 8/– 7–8 (7.6 ± 0.5) 5 –/16 15/15 16/16 –/16 11–20 (15.5 ± 1.8) 47
Incisor–like teeth on upper jaw on premaxilla 42 18–42 (28.3 ± 10.1) 4 19 14 19 13–35 (25.3 ± 5.8) 55
Incisor–like teeth on lower jaw on dentary 39 12–39 (24.8 ± 11.1) 4 17 12 18 12–38 (24.5 ± 6.5) 55
Cheek scales 5/– 6/– 5–6 (5.6 ± 0.5) 5 –/11 11/– 12/– 12/– 9–16 (11.7 ± 1.3) 44
Vertebral number (anterior) AV 10 10 10 (10.0 ± 0.0) 3 10 10 3 10 (10.0 ± 0.0) 27
Vertebral number (caudal) CV 15 15 15 (15.0 ± 0.0) 3 16 16 16 16 (16.0 ± 0.0) 27
Isthmus scale rows 10 10–11 (10.5 ± 0.7) 2 9 8 6 6–13 (9.4 ± 1.6) 41
Interorbital scale rows none none none (0.0 ± 0.0) 5 >14 17 17 19 8–22 (18.2 ± 2.8) 51
Dorsal pterygiophores 20 19 19–20 (19.5 ± 0.7) 2 21 21 21 20–21 (21.0 ± 0.2) 27
Anal pterygiophores 11 11 11 (11.0 ± 0.0) 2 12 12 12 12 (12.0 ± 0.0) 27
Total pterygiophores 31 30 30–31 (30.5 ± 0.7) 2 33 33 33 32–33 (33.0 ± 0.2) 27
…continued on the next page
Zootaxa 3751 (1) © 2013 Magnolia Press · 17
TAXONOMIC REVISION OF KYPHOSIDAE
TABLE 1.
(
continued
)
Kyphosus azureus Kyphosus bigibbus
USNM 39629
ermosilla azurea
(HT)
USNM 87547
H
ermosilla robusta
(HT)
Range of character, for
specimens examined,
including holotype and non-
types.
n
MNHN B–2162
K
yphosus bigibbus
(HT)
SMNS 23084–1
P
imelepterus fallax
(PLT)
SMNS 23084–2
P
imelepterus fallax
(PLT)
MNHN 0000–9601
P
imelepterus bosquii
(HT)
Range of character, for
specimens examined,
including holotype, paratypes
and non-types.
n
Proportional measurements (in %SL)
Head length 28.4 26.6 26.6–33.0 (29.5 ± 2.5) 5 22.1 29.3 27.1 26.6 21.3–30.7 (25.8 ± 2.0) 56
Body width 14.6 13.7–16.2 (14.6 ± 1.1) 4 14.1 12.8 13.2 11.6–26.4 (16.6 ± 2.3) 47
Body depth 50.6 50.3 36.2–50.6 (44.1 ± 6.2) 5 42.2 45.4 43.8 42.9 35.4–49.9 (43.1 ± 3.2) 56
Caudal peduncle length, dorsal 22.2 20.4 20.4–24.1 (21.7 ± 1.5) 5 19.5 18.9 21.9 22.5 17.2–24.1 (20.4 ± 1.9) 55
Caudal peduncle depth 14.9 13.7 13.1–14.9 (13.9 ± 0.7) 5 11.4 12.7 13.1 12.3 9.5–14.1 (11.7 ± 1.0) 56
Snout length 6.0 8.1 6.0–8.1 (7.0 ± 1.1) 5 8.9 7.8 6.5 6.0 5.0–9.0 (6.9 ± 1.0) 55
Eye diameter 7.3 7.0 7.0–11.0 (8.7 ± 1.7) 5 4.4 9.3 8.4 7.5 4.4–9.7 (7.1 ± 1.3) 56
Interorbital width 10.9 10.9–11.9 (11.3 ± 0.4) 4 11.6 11.6 10.3 10.1–13.0 (11.5 ± 0.7) 55
Upper jaw length 7.0 7.0 7.0–9.2 (7.8 ± 1.0) 5 7.4 8.1 6.7 4.8–9.4 (7.2 ± 0.9) 48
Postorbital distance 13.7 13.7 12.7–15.0 (13.8 ± 0.8) 5 8.9 12.2 13.6 12.3 8.9–14.7 (12.3 ± 1.0) 49
Gill slit opening height 21.8 19.6 19.3–21.8 (20.2 ± 1.1) 5 20.6 21.1 19.4 16.5–24.4 (19.9 ± 1.8) 48
Anterior of pectoral fin to anterior of pelvic fin 6.3 3.9 3.9–9.8 (6.8 ± 2.6) 5 5.3 5.1 6.7 4.3–12.5 (8.8 ± 2.2) 44
Isthmus width 4.8 4.0–4.8 (4.5 ± 0.3) 4 3.8 3.4 3.0 3.0–5.2 (3.9 ± 0.4) 48
Preanal length 57.6 63.7 57.6–67.3 (62.1 ± 3.7) 5 61.1 58.2 57.6 60.7 52.5–65.0 (60.3 ± 2.5) 56
Predorsal length 34.0 35.5 34.0–42.6 (37.7 ± 3.5) 5 38.5 33.8 36.1 30.7–41.4 (35.9 ± 2.5) 54
Prepectoral length 26.4 24.5 24.5–32.2 (28.4 ± 3.0) 5 21.1 29.2 26.9 26.7 21.1–31.2 (25.7 ± 2.2) 49
Preventral length 35.6 30.7 30.7–37.5 (34.3 ± 2.5) 5 37.8 34.0 34.2 28.0–43.1 (34.8 ± 3.2) 48
Dorsal fin base length 47.1 44.1 41.0–47.1 (43.5 ± 2.3) 5 51.8 44.9 44.0 45.4 42.2–52.2 (46.6 ± 2.2) 56
Soft dorsal fin base length 15.7 16.8 15.7–21.1 (17.9 ± 2.1) 5 23.5 21.7 19.4 19.5 16.2–25.8 (20.4 ± 2.1) 56
Spinous dorsal fin base length 31.4 27.3 19.9–31.4 (25.6 ± 4.3) 5 28.3 23.3 24.6 25.9 18.5–31.9 (26.3 ± 2.6) 56
1st dorsal fin spine length 3.7 3.0 3.0–5.1 (3.8 ± 0.8) 5 3.7 4.4 5.1 4.2 2.1–5.1 (3.5 ± 0.7) 55
6th dorsal fin spine length 13.8 15.1 13.8–15.1 (14.6 ± 0.5) 5 11.7 14.0 12.4 12.7 9.0–14.5 (12 ± 1.4) 56
4th dorsal fin ray length 8.6 9.4 8.6–13.0 (10.5 ± 1.7) 5 8.5 10.2 11.2 9.1 5.5–13.1 (8.7 ± 1.4) 56
3rd anal fin spine length 7.3 4.1 4.1–8.7 (7.2 ± 1.8) 5 5.9 5.7 8.2 7.1 3.8–9.8 (6.1 ± 1.1) 56
2nd anal fin ray length 13.8 10.2 10.2–16.6 (14.1 ± 2.5) 5 11.4 11.1 11.7 9.7 6.0–13.2 (10.4 ± 1.4) 56
Pectoral fin length 19.6 19.6 19.3–21.9 (20.1 ± 1.0) 5 16.1 19.2 19.1 16.4 13.6–21.1 (17.7 ± 1.5) 54
Anal fin base length 17.5 18.9 14.0–18.9 (17.1 ± 2.0) 5 20.9 18.9 21.1 20.5 18.1–24.8 (20.8 ± 1.5) 56
Caudal fin length 19.4 15.5 15.5–20.5 (18.1 ± 2.0) 5 17.2 19.2 16.6 17.7 12.9–26.7 (19.1 ± 3.2) 56
Pelvic fin length 19.6 16.9 15.2–19.6 (17.4 ± 2.0) 5 16.4 18.0 14.5 8.7–21.3 (17.1 ± 1.9) 55
End of pelvic fin to anal fin 2.4 16.4 2.4–18.5 (10.4 ± 6.8) 5 3.9 5.6 13.4 3.6–14.0 (9.1 ± 2.7) 46
KNUDSEN & CLEMENTS
18 · Zootaxa 3751 (1) © 2013 Magnolia Press
TABLE 2.
Counts and measurements of Kyphosus cinerascens. Values in parentheses indicate mean and standard deviation.
Kyphosus cinerascens
AMS.I 16342–001
P
achymetopon
s
quamosum (HT)
MNHN 0000–8814
Cantharus maculatus
(HT)
MNHN 0000–9603
P
imelepterus
altipinnoides (HT)
MNHN A–1993
K
yphosus cinerascens
(Neo type)
MNHN 0000–9659
P
imelepterus
dussumieri (ST)
MNHN 0000–1313
P
imelepterus
altipinnoides (HT)
MNHN 0000–4784
P
imelepterus
raynaldii (HT)
AMS.I 16349–001
Scorpis vinosa
(HT)
MNHN 0000–0838
P
imelepterus
altipinnoides (PT)
Range of character, for
specimens examined,
including holotype, paratypes
and non-types.
n
Total length (mm) 326 154 282 122 273 93 104 333 64–452 (201.5 ± 86.5) 72
Standard length (mm) 278 31 131 237 101 231 77 86 252 31–382 (167.1 ± 75.4) 74
Counts
Dorsal fin spines XI XI XI XI XI X XI XI XI X–XI (11.0 ± 0.2) 73
Soft dorsal fin rays 12 12 12 12 12 12 12 13 12 12–13 (12.1 ± 0.3) 72
Anal fin spines III III III III III III III III III III (3.0 ± 0.0) 74
Soft anal fin rays 11 11 11 11 11 12 11 11 11 11–13 (11.2 ± 0.5) 69
Pectoral fin rays 18/18 19/18 18/19 18/19 19/19 19/18 18/18 20/19 18/19 16–20 (18.1 ± 0.8) 69
Pelvic fin rays I, 5 I, 5 I, 5 I, 5 I, 5 I, 5 I, 5 I, 5 I, 5 I, 5 (5.0 ± 0.0) 74
Caudal fin rays 17 17 17 17 17 17 17 17 17 16–17 (17.0 ± 0.1) 57
Gill rakers upper limb on first gill arch 8/– 7/7 8/8 7/8 10/10 7/7 8/8 6/6 8/– 6–10 (7.3 ± 0.7) 41
Gill rakers lower limb on first gill arch 19/– 19/19 21/21 20/20 20/20 21/21 20/20 19/19 19/– 17–22 (19.4 ± 0.9) 39
Gill rakers, total on first gill arch 27/– 26/26 29/29 27/28 30/30 28/28 28/28 25/25 27/– 25–30 (26.7 ± 1.1) 40
Pored scales in lateral line 51/53 53/52 57/57 57/58 58/60 55/56 55/53 53/54 57/57 50–60 (54.2 ± 2.3) 61
Pored scales in caudal fin 6/– 5/– 6/– 5/– 6/– 6/– 5/– 5–8 (6.0 ± 0.9) 57
Scale rows above lateral line 11/– 10/– 10/– 11/11 11/ 11/– 10/– 10/– 10/– 9–13 (10.6 ± 0.8) 58
Scale rows below lateral line 19/20 19/18 19/19 20/21 18/19 19/20 19/20 19/19 17/18 17–21 (18.8 ± 0.9) 57
Total scales in lateral line 59/59 65/63 69/67 66/68 73/72 64/63 68/66 59/61 63/65 56–75 (65.1 ± 3.2) 61
Scale rows in longitudinal row 55/54 54/53 61/60 59/55 60/61 57/56 53/54 52/52 58/57 50–61 (55.5 ± 2.4) 27
Postorbital scales 14/15 –/– 15/16 16/18 17/16 16/17 15/15 14/13 17/16 11–18 (14.8 ± 1.5) 56
Incisor–like teeth on upper jaw on premaxilla 33 30 32 18 31 20 21 31 14–39 (28.3 ± 5.5) 49
Incisor–like teeth on lower jaw on dentary 41 26 33 20 34 12 16 32 12–41 (28.3 ± 7.0) 48
Cheek scales 11/– 8/10 10/– –/13 11 13/13 11/– 9/– 13/13 8–14 (10.9 ± 1.2) 56
Vertebral number (anterior) AV 10 10 10 10 10 10 10 10 10 10 (10.0 ± 0.0) 33
Vertebral number (caudal) CV 16 16 16 16 16 16 16 16 16 16 (16.0 ± 0.0) 33
Isthmus scale rows 9 8 10 9 10 9 8 9 8–12 (9.1 ± 0.9) 44
Interorbital scale rows 14 18 14 17 20 17 15 15 13–20 (16.5 ± 1.5) 46
Dorsal pterygiophores 21 21 21 21 21 21 21 20–21 (21.0 ± 0.2) 31
Anal pterygiophores 12 12 12 12 12 12 12 12 (12.0 ± 0.0) 31
Total pterygiophores 33 33 33 33 33 33 33 32–33 (33.0 ± 0.2) 31
…continued on the next page
Zootaxa 3751 (1) © 2013 Magnolia Press · 19
TAXONOMIC REVISION OF KYPHOSIDAE
TABLE 2.
(
continued
)
Kyphosus cinerascens
AMS.I 16342–001
P
achymetopon
s
quamosum (HT)
MNHN 0000–8814
Cantharus maculatus
(HT)
MNHN 0000–9603
P
imelepterus
altipinnoides (HT)
MNHN A–1993
K
yphosus cinerascens
(Neo type)
MNHN 0000–9659
P
imelepterus
dussumieri (ST)
MNHN 0000–1313
P
imelepterus
altipinnoides (HT)
MNHN 0000–4784
P
imelepterus
raynaldii (HT)
AMS.I 16349–001
Scorpis vinosa
(HT)
MNHN 0000–0838
P
imelepterus
altipinnoides (PT)
Range of character, for
specimens examined,
including holotype, paratypes
and non-types.
n
Proportional measurements (in %SL)
Head length 24.3 32.2 28.0 25.6 26.4 24.9 30.4 30.4 26.9 18.0–32.2 (26.8 ± 2.5) 61
Body width 17.8 15.7 15.1 14.2 13.5 16.2 15.2 16.3 17.1 10.3–22.8 (16.2 ± 2.2) 59
Body depth 44.6 38.6 44.0 44.7 45.3 41.8 45.6 46.2 42.4 34.7–47.8 (42.8 ± 2.6) 61
Caudal peduncle length, dorsal 19.5 19.7 24.6 22.9 20.4 23.8 20.6 18.8 23.1 15.1–27.8 (21.9 ± 2.2) 59
Caudal peduncle depth 12.8 14.4 13.3 13.7 13.5 12.2 14.6 13.6 12.8 8.9–14.6 (13.0 ± 0.8) 59
Snout length 7.3 6.3 7.7 8.7 7.1 6.0 7.6 5.2 7.6 3.9–9.1 (6.9 ± 1.2) 61
Eye diameter 6.5 11.4 8.2 7.0 8.0 7.0 9.5 9.2 6.2 5.0–11.4 (7.9 ± 1.2) 61
Interorbital width 9.9 13.9 11.0 10.1 12.2 10.9 12.3 12.8 10.3 7.5–13.9 (11.2 ± 0.9) 61
Upper jaw length 7.1 9.5 8.2 7.7 7.3 7.4 8.7 9.1 7.9 5.8–9.5 (7.9 ± 0.7) 59
Postorbital distance 11.2 12.9 12.8 11.9 12.5 11.5 14.1 13.8 12.5 8.8–15.1 (12.4 ± 1.0) 50
Gill slit opening height 19.2 22.2 21.1 20.8 23.2 19.8 22.9 23.8 21.3 16.4–23.8 (20.1 ± 1.6) 50
Anterior of pectoral fin to anterior of pelvic fin 12.4 5.4 9.4 9.9 7.5 10.6 8.1 12.5 10.3 5.2–13.4 (8.9 ± 2.0) 49
Isthmus width 4.2 3.4 4.0 5.5 4.5 4.6 4.3 4.6 5.0 3.0–11.1 (4.5 ± 1.1) 50
Preanal length 60.4 58.9 58.7 59.1 59.8 59.8 61.5 62.2 59.5 56.6–64.0 (59.6 ± 1.8) 60
Predorsal length 35.9 41.9 38.7 36.5 38.6 39.6 41.1 36.6 33–43.7 (37.3 ± 2.3) 59
Prepectoral length 25.6 32.4 26.8 26.1 27.7 24.6 28.3 30.1 26.0 17.8–34.7 (26.8 ± 2.7) 59
Preventral length 30.6 34.8 35.1 34.2 36.5 33.9 36.3 39.4 34.0 25.0–42.3 (35.0 ± 2.7) 59
Dorsal fin base length 45.0 43.9 42.9 47.0 45.0 40.4 46.4 47.9 46.2 34.4–49.5 (45.2 ± 2.3) 61
Soft dorsal fin base length 19.9 20.2 14.5 21.9 22.4 18.8 17.3 20.9 21.2 14.5–23.0 (19.5 ± 1.8) 61
Spinous dorsal fin base length 25.0 23.7 28.5 26.9 22.6 21.6 29.1 27.0 25.0 18.8–31.8 (25.7 ± 2.3) 61
1st dorsal fin spine length 2.8 3.6 3.1 3.9 1.9 4.1 2.9 3.1 1.7–6.8 (3.2 ± 0.8) 63
6th dorsal fin spine length 9.7 10.3 10.3 11.9 9.5 11.2 10.4 7.9–15.3 (11.0 ± 1.2) 62
4th dorsal fin ray length 15.6 13.9 12.8 13.6 12.8 15.0 15.1 14.3 8.7–19.1 (13.8 ± 1.7) 64
3rd anal fin spine length 4.9 6.4 5.6 4.8 7.0 5.4 5.5 7.0 6.5 4.8–8.4 (6.6 ± 0.9) 65
2nd anal fin ray length 17.2 17.8 16.4 15.7 16.3 15.4 16.7 20.1 10.4–22.3 (16.3 ± 2.1) 64
Pectoral fin length 16.0 20.2 19.7 16.6 18.9 18.6 22.6 20.9 19.4 12.1–22.6 (19.3 ± 1.5) 61
Anal fin base length 19.6 20.2 19 21 22.9 20.3 21.6 21.8 17.3 12.8–23.4 (19.7 ± 1.8) 61
Caudal fin length 17.3 17.6 19.0 20.8 18.2 21.3 21.0 15.5–28.1 (20.9 ± 2.6) 70
Pelvic fin length 16.8 20.5 19.8 18.1 19.0 18.1 20.1 20.6 20.8 11.4–21.4 (19.1 ± 1.5) 65
End of pelvic fin to anal fin 13.1 8.5 7.7 9.1 7.8 8.0 5.3 2.1 4.4 2.5–13.1 (6.1 ± 2.8) 58
KNUDSEN & CLEMENTS
20 · Zootaxa 3751 (1) © 2013 Magnolia Press
TABLE 3.
Counts and measurements of Kyphosus cornelii and Kyphosus elegans. Values in parentheses indicate mean and standard deviation.
K
yphosus cornelii Kyphosus elegans
WAM P.33451–003
K
yphosus cornelii
(Neo type)
Range of character, for specimens
examined, including type and
non-types.
n
ZMB–7071
P
imelepterus elegans
(HT)
MNHN 0000–9818
P
imelepterus
s
andwicensis (HT)
Range of character, for specimens
examined, including holotype and
non-types.
n
Total length (mm) 201 41–406 (253.7 ± 120.2) 26 288 259 189–385 (285.4 ± 51.6) 16
Standard length (mm) 174 33–346 (215.0 ± 104.3) 26 224 220 151–311 (233.9 ± 41.2) 16
Counts
Dorsal fin spines XI XI (11.0 ± 0.0) 26 XI XI X–XI (10.9 ± 0.3) 16
Soft dorsal fin rays 15 14–16 (15.2 ± 0.6) 26 13 13 13–14 (13.1 ± 0.3) 16
Anal fin spines III III (3.0 ± 0.0) 26 III III II–III (2.9 ± 0.3) 16
Soft anal fin rays 15 13–15 (14.2 ± 0.5) 26 12 12 12 (12.0 ± 0.0) 16
Pectoral fin rays 16/– 15–18 (16.5 ± 0.7) 26 18/18 18/18 17–18 (17.8 ± 0.4) 16
Pelvic fin rays I, 5 I, 5 (5.0 ± 0.0) 26 I, 5 I, 5 I, 5 (5.0 ± 0.0) 16
Caudal fin rays 17 17 (17.0 ± 0.0) 20 17 17 17 (17.0 ± 0.0) 16
Gill rakers upper limb on first gill arch 5/– 4–6 (4.7 ± 0.5) 26 7/8 8/8 6–8 (7.2 ± 0.7) 13
Gill rakers lower limb on first gill arch 16/– 13–17 (14.4 ± 0.9) 26 18/19 19/18 16–19 (17.8 ± 0.7) 13
Gill rakers, total on first gill arch 21/– 17–21 (19.1 ± 1.0) 26 26/27 27/26 24–27 (25.3 ± 0.8) 12
Pored scales in lateral line 51/– 41–54 (49.2 ± 3.1) 26 55/55 53/55 52–57 (54.7 ± 1.4) 16
Pored scales in caudal fin 2–5 (3.5 ± 1.0) 13 7/– 6/– 6–8 (7.0 ± 0.8) 16
Scale rows above lateral line 8/– 7–9 (8.0 ± 0.8) 26 13/13 13/12 11–14 (12.4 ± 0.8) 16
Scale rows below lateral line 16/– 13–18 (16.2 ± 1.2) 26 18/19 19/18 17–22 (18.4 ± 1.2) 16
Total scales in lateral line 53/– 50–59 (54.3 ± 2.7) 26 69/66 63/61 61–69 (66.7 ± 2.1) 16
Scale rows in longitudinal row 52/– 45–56 (50.1 ± 2.3) 23 56/54 54/54 52–62 (57.2 ± 2.4) 16
Postorbital scales 5–8 (7.0 ± 1.1) 13 17/16 16/16 12–19 (15.8 ± 1.6) 16
Incisor–like teeth on upper jaw on premaxilla 34 14–41 (30.4 ± 7.1) 25 40 34 30–41 (36.6 ± 3.2) 15
Incisor–like teeth on lower jaw on dentary 28 14–39 (29.8 ± 7.5) 24 37 32 28–41 (36.1 ± 3.3) 15
Cheek scales 3–7 (4.8 ± 1.2) 13 13/– 13/12 10–14 (12.4 ± 1.2) 16
Vertebral number (anterior) AV 10 10 (10.0 ± 0.0) 9 10 10 10 (10.0 ± 0.0) 16
Vertebral number (caudal) CV 15 15 (15.0 ± 0.0) 9 16 16 16 (16.0 ± 0.0) 16
Isthmus scale rows 7–10 (7.8 ± 0.9) 12 10 10 9–12 (9.9 ± 0.8) 16
Interorbital scale rows none none (0.0 ± 0.0) 21 20 18 15–21 (17.9 ± 2.3) 15
Dorsal pterygiophores 24 23–26 (24.3 ± 1.0) 7 22 21 21–22 (21.8 ± 0.4) 16
Anal pterygiophores 16 14–16 (15.1 ± 0.7) 7 13 13 13 (13.0 ± 0.0) 16
Total pterygiophores 40 39–40 (39.4 ± 0.5) 7 35 34 34–35 (34.8 ± 0.4) 16
…continued on the next page
Zootaxa 3751 (1) © 2013 Magnolia Press · 21
TAXONOMIC REVISION OF KYPHOSIDAE
TABLE 3.
(
continued
)
Kyphosus cornelii Kyphosus elegans
WAM P.33451–003
K
yphosus cornelii
(Neo type)
Range of character, for specimens
examined, including type and
non-types.
n
ZMB–7071
P
imelepterus elegans
(HT)
MNHN 0000–9818
P
imelepterus
s
andwicensis (HT)
Range of character, for specimens
examined, including holotype and
non-types.
n
Proportional measurements (in %SL)
Head length 25.9 23.1–33.5 (26.2 ± 2.8) 26 27.1 25.1 24.1–31.6 (26.4 ± 2.1) 16
Body width 13.2–25.6 (16.3 ± 3.1) 13 17.9 12.4 11.0–20.0 (15.5 ± 2.7) 16
Body depth 37.9 33.2–41.9 (37.9 ± 2.2) 26 52.7 48.3 42.9–52.7 (48.1 ± 2.4) 16
Caudal peduncle length, dorsal 23.0 16.6–24.5 (21.5 ± 2.3) 26 21.2 16.7 15.6–21.5 (17.9 ± 1.6) 16
Caudal peduncle depth 10.9 9.0–12.3 (10.8 ± 0.7) 26 13.0 11.0 10.8–13.1 (11.8 ± 0.7) 16
Snout length 8.0 4.8–9.5 (7.5 ± 1.1) 26 6.5 7.9 5.2–8.5 (6.6 ± 0.9) 16
Eye diameter 6.3 4.6–12.3 (6.9 ± 2.2) 26 8.2 7.2 6.6–9.4 (7.8 ± 1.0) 16
Interorbital width 11.5 5.1–12.3 (10.8 ± 1.3) 26 13.1 11.0 11.0–13.8 (12.3 ± 1.0) 16
Upper jaw length 5.3–9.4 (7.0 ± 1.5) 12 7.2 7.1 4.6–9.5 (7.3 ± 1.0) 16
Postorbital distance 11.5–13.9 (12.7 ± 0.7) 13 13.3 11.2 10.7–14.8 (12.8 ± 1.0) 16
Gill slit opening height 15.1–21.4 (17.7 ± 1.9) 13 22.9 20.8 19.1–25.0 (21.7 ± 1.5) 16
Anterior of pectoral fin to anterior of pelvic fin 1.9–10.9 (5.1 ± 2.7) 13 10.4 5.2 4.4–10.9 (7.9 ± 2.1) 16
Isthmus width 3.7–5.4 (4.5 ± 0.5) 12 5.4 5.1 4.6–5.8 (5.2 ± 0.4) 16
Preanal length 52.3 51.7–62.5 (56.7 ± 3.1) 26 63.1 59.9 56.9–63.1 (60.1 ± 2.1) 16
Predorsal length 29.9 27.2–36.6 (30.9 ± 2.7) 26 33.6 34.6 29.1–36.6 (34.0 ± 1.9) 16
Prepectoral length 23.3–32.8 (27.4 ± 3.4) 13 24.9 25.5 23.1–29.9 (25.6 ± 1.8) 16
Preventral length 28.3–39.7 (33.5 ± 3.1) 13 34.1 33.2 32.1–38.4 (34.9 ± 1.7) 16
Dorsal fin base length 51.1 41.5–55.5 (51.4 ± 2.7) 26 55.2 46.4 45.9–55.2 (49.5 ± 2.7) 16
Soft dorsal fin base length 26.4 17.6–29.1 (24.6 ± 2.2) 26 23.7 22.2 19.8–24.0 (21.8 ± 1.4) 16
Spinous dorsal fin base length 24.7 23.7–31.4 (26.7 ± 2.1) 26 31.4 24.2 24.2–31.4 (27.8 ± 2.0) 16
1st dorsal fin spine length 4.0 2.9–8.2 (4.4 ± 1.6) 23 3.5 3.8 2.3–4.5 (3.4 ± 0.5) 16
6th dorsal fin spine length 10.3 7.1–18.9 (10.7 ± 3.1) 25 12.5 12.2 9.8–13.6 (11.8 ± 1.1) 16
4th dorsal fin ray length 8.6 6.1–14.1 (8.5 ± 2.0) 26 10.9 12.0 7.7–12.0 (9.7 ± 1.2) 16
3rd anal fin spine length 7.5 4.3–11.2 (6.9 ± 1.8) 26 6.2 6.1 3.8–8.1 (5.9 ± 1.0) 15
2nd anal fin ray length 12.1 9.5–16.7 (11.9 ± 1.9) 26 13.0 12.4 9.9–15.8 (12.5 ± 1.8) 16
Pectoral fin length 17.8 15.0–22.9 (17.8 ± 2.2) 26 20.6 19.3 16.5–21.3 (19.7 ± 1.2) 16
Anal fin base length 28.7 22.2–28.7 (25.3 ± 1.9) 26 24.9 21.5 20.4–25.7 (22.7 ± 1.6) 16
Caudal fin length 15.5 15.4–24.3 (18.2 ± 2.5) 20 28.9 17.7 15.3–28.9 (22.0 ± 4.1) 16
Pelvic fin length 15.5 12.5–21.4 (16.1 ± 2.4) 26 19.8 17.5 15.0–19.8 (18.0 ± 1.5) 16
End of pelvic fin to anal fin 3.3–13.5 (6.4 ± 3.1) 13 7.0 11.9 2.4–13.1 (8.8 ± 3.3) 16
KNUDSEN & CLEMENTS
22 · Zootaxa 3751 (1) © 2013 Magnolia Press
Kyphosus bigibbus Lacepède, 1801
(Common name: darkfin drummer, brown chub, gray chub, gray rudderfish)
(Figure 2, 3, 5, 6, Table 3, 8–18)
Kyphosus bigibbus Lacepède 1801: p. 114, 115 (type locality: Ile de France, Madagascar), holotype: MNHN B-2162 [dried
skin]; Sonnini 1803: p. 124, 125; Fowler 1931: p. 248 (Port Sudan, Red Sea); Fowler 1941: p. 252; Bauchot 1963: p. 174;
Zama 1976: p. 100, fig. 1 (not K. bigibbus, but K. sectatrix); Araga 1984: p. 166, pl. 152-D, E; Dor 1984: p. 166; Smith
1986: p. 603, fig. 189.1 (key) (South Africa); Pequeño 1989: p. 62; Paulin et al. 1989: p. 194, fig. 127.4b; Nakabo 1993: p.
773 (key, Southern Japan); Kuiter 1993: p. 212 (photo is showing K. sectatrix); Francis 1993: p. 162 (probably mistaken K.
sectatrix for K. bigibbus); Goren and Dor 1994: p. 46; Randall 1995: p. 246, fig. 642; Clements 1997: p. 220, fig. 1F
(photo is of a K. sectatrix); Allen 1997: p. 144, pl. 53, fig. 12 (tropical Australia and Southeast Asia); Anderson et al. 1998:
p. 25; Myers 1999: p. 173, pl. 91-A–B (both photos are of K. sectatrix); Fricke 1999: p. 322 (Reunion, Mauritius); Johnson
1999: p. 738 (probably mistaken K. sectatrix for K. bigibbus); Randall 1999: p. 17, BPBM 16712, BPBM 16736, BPBM
16835 (2), (probably mistaken K. sectatrix for K. bigibbus); Allen 2000: p. 105; Kuiter 2000: p. 212 (photo is showing K.
sectatrix); Pereira 2000: p. 11 (Mozambique); Randall and Earle 2000: p. 15; Randall and Lim 2000: p. 623; Francis 2001:
p. 49, pl. 72 (photo is of a K. sectatrix); Hutchins 2001a: p. 36; Hutchins 2001b: p. 264, app. 1; Sakai 2001: p. 3292 (Indo-
west Pacific, Red Sea, Australia, Japan); Nakabo 2002: p. 958 (key, southern Japan); Allen and Adrim 2003: p. 41 (Papau
to Sumatra); Clements 2003: p. 130; Manilo and Bogorodsky 2003: p. S111 (Indian Ocean, Arabian Sea); Myers and
Donaldson 2003: p. 627; Letourneur et al. 2004: p. 212 (Reunion Island); Lobel and Lobel 2004: p. 72 (BPBM 4043,
BPBM 4044); Sakai and Nakabo 2004: p. 25, fig. 4, 5, table 3; Heemstra et al. 2004: p. 3321, table 1; Heemstra and
Heemstra 2004: p. 250 (Indian Ocean, Pacific Ocean, Red Sea); Pequeño 2004: p. 96 (Pacific Ocean, Salas Y Gomez)
(probably mistaken K. sectatrix for K. bigibbus); Randall 2005: p. 305 (Indian Ocean, Pacific Ocean); Hoese and Bray
2006: p. 1322 (Pacific Ocean, Indo- west Pacific Ocean); Sakai and Nakabo 2006: p. 354 (key); van der Elst 1989: p. 224
(eastern South Africa); Fricke et al. 2009: p. 70 (Reunion Island); Golani and Bogorodsky 2010: p. 36 (Red Sea);
Motomura et al. 2010: p. 133, fig. 257; Beets et al. 2010: p. 41; Knudsen and Clements 2013: p. 1; Nemeth and Kadison
2013: p. 2.
Pimelepterus bosquii Lacepède 1802: p. 429, pl. ix, (type locality: North Atlantic, South Carolina), holotype: MNHN 0000-
9601 (1), Eschmeyer (2013) incorrectly lists MNHN 0000-2977 (1) as a second syntype for Pimelepterus bosquii, but
Bauchot (1963) and the MNHN catalogue list only MNHN 0000-9601 as holotype; Tortonese 1935: p. 242 (regarded as
junior synonym of Kyphosus sectatrix); Bauchot 1963: p. 171–172.
Dorsuarius nigrescens Lacepède 1803: p. 482, 483 (No type locality, but likely Fort Dauphin, Madagascar), holotype: MNHN
B-2162 [dried skin]; Bauchot 1963: p. 169 (regarded as synonym of K. bigibbus); Fricke 1999: p. 322; Sakai and Nakabo
2004: p. 25 (recognised as synonym of K. bigibbus).
Kyphosus fuscus Lacepède 1803: p. 484–485 (type locality: Ile de France, Madagascar) described as Xyster fuscus based on the
description by Philibert Commerçon (unpubl. manuscript), type: MNHN B-2162 [dried skin]; redescribed as Pimelepterus
fuscus by Cuvier in Cuvier and Valenciennes 1831: p. 264; Jordan and Evermann 1905: p. 248 (Honolulu); McCulloch and
Waite 1916: p. 442; Jordan and Jordan 1922: p. 51; Fowler 1925: p. 27; Fowler and Ball 1925: p. 15; Fowler 1927: p. 15;
Fowler 1928: p. 222; Fowler 1929: p. 645; Allen et al. 1976: p. 406 (Lord Howe Island, Norfolk Island).
Xyster fuscus Lacepède 1803: p. 484, 485 (type locality: Ile de France, Madagascar) based on the description by Philibert
Commerçon (unpubl. manuscript), type: MNHN B-2162 [dried skin], objective synonym of K. bigibbus; redescribed as
Pimelepterus fuscus by Cuvier in Cuvier and Valenciennes 1831: p. 264; Fowler 1933: p. 205; Bauchot 1963: p. 169, 174
(
regarded as synonym of K. bigibbus); Dor 1984: p. 167; Fricke 1999: p. 323; Sakai and Nakabo 2004: p. 25 (recognised
as synonym of K. bigibbus).
Xyster nigrescens described as Dorsuarius nigrescens by Lacepède 1803: p. 482, 483, redescribed as Xyster nigrescens by
Cuvier in Cuvier and Valenciennes 1831: p. 264; Fowler 1933: p. 211 (regarded as synonym of K. bigibbus).
Cyphosus fuscus Gilchrist and Thompson 1908: p. 174 (Durban); Regan 1908: p. 245 (Kosi Bay); Gilchrist and Thompson
1917: p. 324; Fowler 1933: p. 211 (regarded as synonym of K. bigibbus).
Pimelepterus fuscus redescribed from Xyster fuscus by Cuvier in Cuvier and Valenciennes 1831: p. 264; Rüppell 1835: p. 34,
pl. 10, fig. 3 (Red Sea); Pappe 1853: p. 23 (Cape of Good Hope); Günther 1859: p. 498 (Red Sea); Bleeker 1860: p. 53;
Castelnau 1861: p. 34; Martens 1866: p. 378 (Kosier, Red Sea); Günther 1874: p. 68 (Tahiti and Hawaii); Day 1875: p.
143; Peters 1876: p. 438 (Mauritius); Klunzinger 1884: p. 65 (Red Sea); Günther 1889: p. 6 (Cape of Good Hope);
Boulenger 1892: p. 134; Steindachner 1900: p. 489 (Honolulu, Laysan); Zugmayer 1913: p. 12 (Oman); Fowler 1933: p.
206 (regarded as synonym of K. bigibbus).
Pimelepterus boscii described as Pimelepterus bosquii by Lacepède 1802, spelled as Pimelepterus boscii by Cuvier and
Valenciennes 1831 p. 258, pl. 187, (type locality: North Atlantic, South Carolina); recognised as Pimelepterus boscii
variant sicula by Döderlein 1883 based on Pimelepterus bosquii Lacepède 1802 and Pimelepterus boscii by Cuvier and
Valenciennes 1831; DeKay 1842: p. 100, pl. xx (fig. 56); Günther 1859: p. 497; Peters 1876: p. 246; Bean 1880: p. 94;
Goode and Bean 1882: p. 238; Desoutter 1973: p. 420; Tortonese 1986: p. 913; Desoutter 1990: p. 831.
Pimelopterus fuscus spelled as ‘Pimelopterus’ by Klunzinger 1870: p. 796; Fowler 1933: p. 206 (regarded as synonym of K.
bigibbus).
Pimelepterus bosci Jordan and Gilbert 1879: p. 378; Jordan and Gilbert 1882: p. 561.
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TAXONOMIC REVISION OF KYPHOSIDAE
Pimelepterus Boscii sicula variant Döderlein 1883: p. 83 (type-locality: Gulf of Palermo, Sicily, Italy), no types known;
Döderlein 1891: p. 234; Vinciguerra 1893: p. 312; Desoutter 1973: p. 420; Desoutter 1990: p. 831.
Cyphosus bosqui Jordan 1884: p. 128 (Key West, Florida).
Pimelepterus fallax Klunzinger 1884: p. 64 (type locality: Al-Qusair, Red Sea, Egypt), lectotype: SMNS 3495, paralectotypes:
SMNS 23084–23085 [except 3495] (2), based on Pimelopterus tahmel (synonym of K. cinerascens) of Klunzinger 1870:
p. 795; Fowler 1933: p. 211 (regarded as synonym of K. bigibbus); Dor 1984: p. 167; Fricke 1992: p. 11; Fricke 2005: p.
43; Sakai and Nakabo 2004: p. 25 (recognised as synonym of K. bigibbus).
Pimelepterus Bosci Ariola 1904: p 165.
Cyphosus boscii Murray and Hjort 1912: p. 614.
Kyphosus gibsoni Russell 1983a: p. 66 (K. gibsoni is likely to have been misidentified K. bigibbus); Kuiter 1993: p. 211 (two
photos of K. gibsoni are photos of K. bigibbus) [K. gibsoni was recognised as a junior synonym of K. vaigiensis by Sakai
and Nakabo (1995)]; Kuiter 2000: p. 211 (two photos of K. gibsoni are photos of K. bigibbus); Hoese and Bray 2006: p.
1317, p. 1323 (likely to have been confused with K. bigibbus).
Kyphosus sydneyanus Kuiter 1993: p. 211 (photo of east coast form of K. sydneyanus is a photo of K. bigibbus); Kuiter 2000: p.
211 (photo of east coast form of K. sydneyanus is a photo of K. bigibbus).
Kyphosus sectator Canas et al. 2005: p. 1536, fig 1A (photo of Algarve specimen is of K. bigibbus).
Material examined
61 specimens (62-527 mm SL): AIM 6496, 213 mm, Three Kings Islands, North New Zealand; AIM MA30184,
425
mm, 36°07' S, 175°29' E; AIM MA30189, 219 mm, 36°07' S, 175°29' E; AMS.I 14060, 105 mm, 31°31' S,
159°05' E; AMS.I 14061, 106 mm, 31°31' S, 159°05' E; AMS.I 17373-013, 253 mm, 31°32' S, 159°04' E; AMS.I
22455-001, 235 mm, 29°05' S, 168°00' E; AMS.I 38651-001, 85 mm, 32°32' S, 152°28' E; AMS.I 44114-002, 182
mm, 33°54' S, 151°15' E; AMS.I 7852, 160 mm, 31°31' S, 152°57' E; AMS.IB 6367, 251 mm, 19°21' S, 146°57' E;
BMNH 1845.10.29.47, 451 mm, Red Sea; BMNH 1845.6.22, 417 mm; BMNH 1864.4.22.16, 188 mm, Madeira;
BMNH 1873.4.3.3, 219 mm, Tahiti; BMNH 1891.2.9.21, 477 mm, 23°61' N, 58°45' E; BMNH 1905.3.17.30-1, 211
mm, Jamaica; BMNH 1941.4.18.30, 262 mm, Mauritius, Pte-aux sables; BMNH 1953.11.169, 194 mm, 32°39' N,
16°55' W; BMNH 1960.3.15.913, 165 mm, Sudanese, Red Sea, Khor, Inkeifail; BMNH 2010.8.4.3, 76 mm;
MNHN 0000-9601 (holotype of Pimelepterus bosquii), 123 mm, 30°00' N, 75°00' W; MNHN 0000-9626, 109 mm,
20°00' N, 39°00' E; MNHN 1975-0380, 368 mm, 17°58' S, 38°37' W; MNHN 1977-0936, 127 mm, 29°33' N,
34°57' E; MNHN A-0764, 427 mm, 34°21' S, 18°25' E; MNHN A-2237, 270 mm, 15°58' S, 38°28' W; MNHN B-
2162 (holotype of Kyphosus bigibbus), >298 mm, 20°00' S, 42°30' E; NMNZ P.005848, 198 mm, 35°30' S, 174°44'
E; NMNZ P.017222, 100 mm, 35°00' S, 174°15' E; NMNZ P.018478, 139 mm, 34°08' S, 172°09' E; NMNZ
P.034930, 176 mm, 33°25' S, 179°00' E; NMNZ P.041290, 527 mm, 29°10' S, 177°50' W; NMNZ P.045653, 294
mm, 36°26' S, 175°47' E; NMNZ P.045657, 297 mm, 36°04' S, 175°20' E; SMNS 23084 1 of 2 (paralectotype of
Pimelepterus fallax), 102 mm, 26°06' N, 34°17' E; SMNS 23084 2 of 2 (paralectotype of Pimelepterus fallax), 91
mm, 26°06' N, 34°17' E; WAM P.22669-001, 202 mm, 22°41' S, 113°41' E; WAM P.23949, 317 mm, 20°29' S,
116°32' E; WAM P.25117-002, 240 mm, 20°28' S, 116°36' E; WAM P.25743-004, 152 mm, 32°01' S, 115°28' E;
WAM P.26665-001, 153 mm, 26°08' S, 113°10' E; WAM P.29122-001 (2 specimens), 136–153 mm, 20°25' S,
115°32' E; WAM P.29335-001, 264 mm, 22°20' S, 113°51' E; WAM P.33451-001, 270 mm, 28°41' S, 113°49' E;
WAM P.4600, 273 mm, 24°29' S, 113°25' E; ZMB-11429, 164 mm, Bermudas; ZMB-7866, 100 mm, 26°06' N,
34°16' E; ZMB-7997, 186 mm, 26°06' N, 34°16' E; ZMUC journ. 1, 140 mm, Nagasaki, J. Jordan; ZMUC journ.
254, 62 mm, Atlantic, Saint Croix; ZMUC journ. 256, 150 mm, Atlantic, Bermuda; ZMUC P518, 75 mm, 23°41'
N, 65°44' W; ZMUC P519, 73 mm, 23°41' N, 65°44' W; no voucher (2 specimens photographed), 260–379 mm,
23°12' S, 113°45' E; no voucher, specimen photographed, 169 mm, 23°01' S, 113°46' E; no voucher (3 specimens
photographed), 268–349 mm, 28°29' S, 113°44' E.
Diagnosis
Terminal mouth, almost in an oblique angle. Scales present in interorbital region. Dorsal fin with XI spines and 11–
1
2 rays. Anal fin with III rays and 10–12 rays. Gill rakers externally on first arch: 4–7 and 13–17 on upper and
lower limb, respectively. Uniform grey or silvery body with weak indications of coloured horizontal scale rows.
Unpaired fins dusky. Pectoral fin relatively short (13.6–21.3 %SL). Anal-fin base relatively long (18.1–24.8 %SL).
Lateral line with 61–76 scales in total, of which 51–60 has pores. Longitudinal row with 54–67 scale rows. Cheek
scales 9–16. Precaudal vertebrae 10, caudal vertebrae 16. Low number of dorsal and anal pterygiophores 20–21
and 12, respectively.
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24 · Zootaxa 3751 (1) © 2013 Magnolia Press
FIGURE 5. Map showing locations for samples and museum material of K. bigibbus used for this study. Museum specimens (black circles), tissue samples (white circles), museum
specimen with tissue sample analysed (white squares) and museum type-specimens (red triangles).
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TAXONOMIC REVISION OF KYPHOSIDAE
FIGURE 6. Ordination plots from principal component analysis for comparison of multiple morphometric and meristic
characters for: A) Kyphosus sectatrix (squares/blue area) and K. bigibbus (triangles/yellow area), type specimens of K. bigibbus
(closed triangles), specimens of K. bigibbus with molecular identity confirmed (open downpointing triangle), type specimens
of K. sectatrix (closed squares), specimens of K. sectatrix with molecular identity confirmed (closed diamonds), B) Kyphosus
sydneyanus (circles/red area) and K. bigibbus (triangles/yellow area), type specimens of K. bigibbus (closed squares),
specimens of K. bigibbus with molecular identity confirmed (closed triangles), type specimens of K. sydneyanus (open
diamond), specimens of K. sydneyanus with molecular identity confirmed (closed circles). Type specimens are noted with
catalog numbers, see text for further details on morphometric and meristic characters used in combination.
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26 · Zootaxa 3751 (1) © 2013 Magnolia Press
Description
Type material.
Holotype (MNHN B-2162, dry skin, right side): D: XI, 12; A: III,
11; P1: –/18; P2: –; PScLL: –/56; TScLL: –/74;
ScRLR: –/66; GRU: –; GRL: –; GRT: –; PCV: –; CV: –; D-Ptg: –; A-Ptg: –.
Paralectotype of Pimelepterus fallax (SMNS 23084 1 of 2): D: XI, 12; A: III, 11; P1: 19/19; P2: I, 5; PScLL: 59/60;
TScLL: 65/67; ScRLR: 57/59; GRU: 6/5; GRL: 14/14; GRT: 20/19; PCV: 10; CV: 16; D-Ptg: 21, A-Ptg: 12.
Paralectotype of Pimelepterus fallax (SMNS 23084 2 of 2): D: XI, 12; A: III, 11; P1: 19/19; P2: I, 5; PScLL: 56/54;
TScLL: 67/66; ScRLR: 64/63; GRU: 4/4; GRL: 14/14; GRT: 18/18; PCV: 10; CV: 16, D-Ptg: 21, A-Ptg: 12.
Holotype of Pimelepterus boscquii (MNHN 0000-9601): D: XI, 12; A: III, 11; P1: –/19; P2: I, 5; PScLL: 58/58;
TScLL: 72/74; ScRLR: 66/65; GRU: 6/6; GRL: 15/15; GRT: 21/21; PCV: 10; CV: 16, D-Ptg: 21, A-Ptg: 12.
Other material (including type material): D: XI, 11–12; A: III, 10–12; P1: 16–20; P2: I, 5; PScLL: 51–60; TScLL:
61–76; ScRLR: 54–67; GRU: 4–7; GRL: 13–17; GRT: 18–22; PCV: 10; CV: 16; D-Ptg: 20–21, A-Ptg: 12.
Body oblong, elliptical in lateral view. Caudal fin not deeply emarginated. Scales ctenoid, present on
i
nterorbital region, and covering postorbital region, cheek and operculum, and arranged in rows along the body
from the posterior edge of the operculum to the caudal fin. Mouth terminal and ventrally oblique. Teeth lanceolate
or incisiform. Small patch of pointed cone-shaped teeth arranged in 3–4 rows on mouth floor and mouth roof
situated well behind outer front row. Head profile slightly pointed, but with a distinct bump on the front of the head
when viewed in profile. Head moderately short (21.3–30.7 %SL), with relative narrow interorbital region (10.1–
13.0 %SL). Snout (5.0–9.0 %SL) as long as eye diameter (4.4–9.7 %SL). Body relatively deep (35.4–49.9 %SL)
and not wide (11.6–26.4 %SL). Caudal peduncle short (17.2–24.1 %SL) and shallow (9.5–14.1 %SL). Anal-fin
base relatively long (18.1–24.8 %SL), but not as long as dorsal-fin base (42.2–52.2 %SL). Spinous part of dorsal
fin (sixth dorsal fin spine 9.0–14.5 %SL) almost as high as the soft-rayed part of dorsal fin (fourth dorsal fin ray
5.5–13.1 %SL). Spinous part of dorsal fin considerably longer (18.5–31.9 %SL) than soft-rayed part of dorsal fin
(16.2–25.8 %SL). Posterior edges of soft-rayed part of anal- and dorsal fin rounded. Pectoral fin moderately long
(13.6–21.1 %SL) with rounded posterior edge. Preanal length moderately short (52.5–65.0 %SL). Height of anal
fin relatively low (second anal fin ray length 6.0–13.2 %SL). Predorsal length relatively long (30.7–41.4 %SL).
Colour when freshV
ariable brownish to silvery with dusky unpaired fins when fresh, which probably is one
of the reasons that specimens of K. bigibbus have been confused with K. sydneyanus. Kyphosus sectatrix can have
similar body colouration, but lacks dusky unpaired fins. The soft part of the dorsal- and the soft part of the anal fin
in K. bigibbus can appear dusky and have black or darker edges. The caudal fin can also appear dusky. The dark
edges on dorsal-, anal- and caudal fin appear pronounced in more adult individuals, but also appear to vary from
habitat to habitat. Although a dark patch occasionally can be present in the posterior-ventral corner of the pectoral-
fin base, this does not appear to be a consistent colour marking and should not be used for identification. The
ventral part of the body more silvery and the dorsal part of the body with a more bronzed to green or dark green
colouration. Cheek and area below eye usually with a white or silvery streak.
Colour of preserved specimens
Often faded to a more uniform brown or yellow colour, or tanned or light
brown. The dusky unpaired fins may not retain their dark colour, and the dark spot on the pectoral fin and the white
streak below the eye on the cheek might also fade and make the fish attain a uniform colour.
Distribution
Previous literature on the distribution of K
. bigibbus is problematic, as specimens from both the East Atlantic and
the West Atlantic have been confused with K. sectatrix. Our recently obtained molecular data from Midwestern
Atlantic kyphosid samples demonstrate the presence of K. bigibbus at San Blas in Panama and at Saint Helena
Island (published separately), and thus K. bigibbus is broadly distributed in the Atlantic Ocean as it is in the Indo-
Pacific region (Figure 5). In Western Australia found from Rottnest Island northwards, and in eastern Australia in
southern part of Queensland and New South Wales south to Montague Island. Found in north-eastern New Zealand
and around the periphery of the Coral Sea at New Caledonia, Lord Howe Island, Norfolk Island and the Kermadec
Islands. Not recorded in the northen part of the Indian Ocean, but likely to be distributed from Burma, along the
coast of India, Pakistan, Iran and Yemen to the Red Sea. Present in the Red Sea (Sakai & Nakabo 2004) and the
southern part of the Arabian Gulf (Figure 5), likely to also be present in the Gulf of Aden and on the coast of
Somalia and south along the East coast of Africa to Madagascar. Present at Madagascar and Reunion Island, and
likely to be present around the Cape of Good Hope. Known from the Mid-Atlantic at Saint Helena, Ascension
Island and the Caribbean Sea on the coast of Panama and Belize (Figure 5). Probably also present along the
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TAXONOMIC REVISION OF KYPHOSIDAE
western coast of Africa from South Africa to São Tomé and Principe. Perhaps also along the North-West coast of
Africa and north towards Portugal. The individual reported by Canas et al. (2005) from the Algarve coast was
identified as K. sectator (i.e. older name K. sectatrix), but the specimen on the photo (Canas et al. 2005, fig. 1A)
resembles K. bigibbus. Recorded at Madeira in the East Atlantic Ocean between the Azores and the Canary Islands
(Figure 5), but not known from the Mediterranean. There is a chance that the records of K. sectatrix at the Canary
Islands (Dooley et al. 1985) and the Azores (Santos et al. 1997) could have included the occasional K. bigibbus. In
the Pacific Ocean known from the coast of Southern Japan, Korea and East China and southwards to Taiwan
(Figure 5) (Sakai & Nakabo 2004), but probably absent from the equator in the Indo-Pacific Ocean. Not recorded
in the South China Sea, and appear absent from Indonesia, Malaysia, Papa New Guinea and Northern Australia.
Most eastern record in the Pacific Ocean is from the Kermadec Islands, north of New Zealand. The single specimen
from Rapa Island listed by Sakai and Nakabo (2004) is likely to be a specimen of K. sectatrix. However, the
distribution appears to be antitropical in the Pacific Ocean, with K. bigibbus being absent from the equator in the
Indo-Pacific Ocean, but present in both hemispheres in subtropical seas, and occasionally also in temperate seas. In
the Atlantic Ocean K. bigibbus is also found at lower latitudes in the Caribbean Sea (Figure 5). Often encountered
as solitary individuals or together with schools of K. sectatrix in the Atlantic and Indo-Pacific, and together with K.
sydneyanus in the south west Pacific. In north-eastern and north-western Australia K. bigibbus form large schools
of hundreds of individuals on rocky and coral reefs. At low latitude reefs K. bigibbus also associates in schools
with K. cinerascens, K sectatrix and K. vaigiensis. Their diet is comprised of a wide mix of green, brown and red
algae. They are found in shallow waters from the surface down to 25 m depth.
Remarks
Eschmeyer (2013) lists P
imelepterus bosquii (Lacepède 1802) as having two existing syntypes (MNHN 0000-9601
and MNHN 0000-2977), however Bauchot (1963), Desoutter (1990) and the MNHN catalogue (available through
the MNHN internet webpage) list only MNHN 0000-9601 as the holotype for Pimelepterus bosquii. MNHN 0000-
9601 and MNHN 0000-2977 are found in the separate jars in MNHN, and labels on the jars indicate that MNHN
0000-9601 is the holotype for Pimelepterus bosquii, and MNHN 0000-2977 is the holotype for Kyphosus sectatrix.
MNHN 0000-2977 was caught in 1768 in the Atlantic Ocean on James Cook’s first voyage (Bauchot 1963,
Wheeler 1986) and brought back to Europe in 1771. This specimen was thus available for the 13th edition of
Systema Natura (Gmelin 1789) to match the description of Perca sectatrix by Linnaeus (1766) that was based on
the drawing presented by Catesby (1743). MNHN 0000-9601 was caught on the East Atlantic coast of North
America (Bauchot 1963), perhaps collected by Louis Augustin Guillaume Bosc between 1796 and 1798, and
brought back to Paris and added to the personal collection of Pierre Marie Auguste Broussonet, who at the time
already held a large part of the ichthyological material collected on the Cook voyages (Bauchot 1969, Whitehead
1969, 1978). This specimen was thus available for Lacepède (1802) to examine for the description of Pimelepterus
bosquii. We consider MNHN 0000-2977 to be the original holotype for Perca sectatrix, collected on James Cook’s
first voyage and examined for the 13th edition of Systema Natura (Gmelin 1789), to which Lacepède (1802)
subsequently added MNHN 0000-9601 as the holotype for Pimelepterus bosquii. We found that MNHN 0000-9601
matches the variation found in K. bigibbus (Figure 6). The low gill raker count and low second anal fin ray make it
certain that this specimen is K. bigibbus. This is not surprising, as K. bigibbus, K. pacificus and K. hawaiiensis in
the Indo-Pacific were considered synonymous until Sakai and Nakabo (2004) correctly split K. bigibbus into three
species.
The finding of K. bigibbu
s in the Atlantic Ocean indicates that the species is more widely distributed than
expected. The Atlantic Ocean was previously considered to contain two endemic species of Kyphosus (K. sectatrix
and K. incisor). It now appears that confounding K. bigibbus and K. sectatrix has not only been a problem in the
Indo-Pacific, but also caused difficulties for correct identification in the Atlantic Ocean. The current presumed
syntype specimen for K. sectatrix (MNHN 0000-9601, originally identified as Pimelepterus bosquii) has a total gill
raker count (21/21) which is lower than the range found in K. sectatrix (22–25) (as defined by MNHN 0000-2977).
We thus consider MNHN 0000-9601 (the holotype for Pimelepterus bosquii) to represent K. bigibbus, and
accordingly Pimelepterus bosquii (Lacepède 1802) becomes a junior synonym of Kyphosus bigibbus (Lacepède
1801).
Sakai and Nakabo (2004) identified Pi
melepterus fallax (SMNS 23084) and Pimelepterus fuscus (BMNH
1845.10.29.47 and MNHN A-0764) as junior synonyms of K. bigibbus. These three specimens have 12 soft dorsal
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28 · Zootaxa 3751 (1) © 2013 Magnolia Press
fin rays, 11–12 soft anal fin rays, more than 65 total scale rows along the lateral line, and SMNS 23084 have fewer
than 23 gill rakers in total on the first gill arch, and thus Pimelepterus fallax and Pimelepterus fuscus match the
variation recorded for K. bigibbus. Also, Sakai and Nakabo (2004) considered the type specimen for K. bigibbus to
be distinct from K. pacificus and K. hawaiiensis based on the length of the second anal fin ray and the catch
locality. With K. sectatrix here considered a junior synonym of K. pacificus (see below), the distributions of K.
bigibbus and K. pacificus are now known to overlap, and the type locality (Mauritius) cannot be used to determine
that the identity of MNHN B-2162 is K. bigibbus. However, an ordination plot m