Zootaxa 1671: 3–31 (2008)                                                                                                  ISSN 1175-5326 (print edition)

www.mapress.com/zootaxa/                                            ZOOTAXA

Copyright © 2008 Magnolia Press

ISSN 1175-5334 (online edition)

Five new species of the damselfish genus Chromis (Perciformes: Labroidei:
Pomacentridae) from deep coral reefs in the tropical western Pacific

RICHARD L. PYLE*, JOHN L. EARLE† & BRIAN D. GREENE‡

Department of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817-2704, USA.1
E-mail: *deepreef@bishopmuseum.org, †earlej001@hawaii.rr.com, ‡bgreene@hawaii.edu.

Table of contents

Abstract ........................................................................................ 3

Introduction ..................................................................................... 3

Material and methods ............................................................................. 4

Chromis abyssus, new species ....................................................................... 6

Chromis brevirostris, new species ................................................................... 10

Chromis circumaurea, new species .................................................................. 15

Chromis degruyi, new species ...................................................................... 18

Chromis earina, new species ....................................................................... 21

Acknowledgements .............................................................................. 25

References ..................................................................................... 26

Appendix: Embedded external hyperlinks ............................................................ 29

Abstract

Five new species of the damselfish genus Chromis2 (Perciformes3: Labroidei4: Pomacentridae5) are described from speci-
mens collected from deep (>60 m) coral-reef habitat in the western Pacific by divers using mixed-gas closed-circuit
rebreather gear. Two of the five new species (C. abyssus and C. circumaurea) are each described from specimens taken at
a single locality within the Caroline Islands (Palau and Yap, respectively); one (C. degruyi) is described from specimens
collected or observed throughout the Caroline Islands, and two (C. brevirostris and C. earina) are described from speci-
mens collected from several localities throughout the Caroline Islands, Fiji, and Vanuatu. All five species can easily be
distinguished from other known Chromis, and from each other, on the basis of color and morphology. These new species
represent the first five scientific names prospectively registered in the official ICZN ZooBank registry6. Moreover, the
electronic online edition of this document has been specially formatted with many embedded links to additional
resources available online via the internet to enhance access to taxonomically-relevant information, and as a demonstra-
tion of the utility of international standards for biodiversity informatics.

Key words: Chromis abyssus, Chromis brevirostris, Chromis circumaurea, Chromis degruyi, Chromis earina, new spe-
cies, taxonomy, cybertaxonomy, mesophotic, mixed-gas diving, rebreather, ZooBank

Introduction

Over the past two decades, the authors have developed and refined techniques for using advanced dive gear
and various gas mixtures to allow safe excursions to depths of 50–150 m, which exceeds the depths that can
be safely reached using conventional SCUBA. Initial exploratory deep dives by the authors at various locali-
ties throughout the tropical Pacific have revealed a rich diversity of coral-reef-associated fishes inhabiting

Accepted by L. A. Rocha: 11 Dec. 2007; published: 1 Jan. 2008                                                                  3

urn:lsid:zoobank.org:pub:68376390-7809-46FF-9EC4-1371B4AAD0FF

these depths (Pyle 1996a, 1996b, 1999, 2000). In addition to the discovery of many new species of fishes, ini-
tial exploration of deep coral reefs (sometimes referred to as “mesophotic” reefs) has led to new insights con-
cerning ecological and biogeographic processes in both the Pacific (Pyle 2000, unpublished data) and the
tropical Atlantic (Feitoza et al. 2005).

The pomacentrid genus Chromis Cuvier 1814 (type species Sparus7 chromis8 Linnaeus 1758) is the largest
genus of the family, with 86 valid species. Fishes of this genus are among the few pomacentrid species known
to inhabit deep coral reefs, perhaps a reflection of their principal food (zooplankton), in contrast to species of
genera that are dependant primarily on benthic algae. At least 34 species of Chromis are known to inhabit
depths of 50 m or more, but only nine are restricted to such depths, and only five (C. okamurai9 Yamakawa
and Randall 1989, C. mirationis10 Tanaka 1917, C. struhsakeri11 Randall and Swerdloff 1973, C. abyssicola12
Allen and Randall 1985, and C. onumai13 Senou and Kudo 2007) are known only from depths of 60 m or
more. Except for two specimens of C. brevirostris collected at a depth of 60 m, all specimens of the new spe-
cies described herein were collected at depths of 85 m or more.

Whereas the print (paper) edition of this article represents the publication through which these five new
scientific names are made available under the current (4th Edition) International Commission of Zoological
Nomenclature (ICZN)14 Code of Nomenclature15, the concurrently distributed electronic online edition is
intended to exemplify the utility of modern international biodiversity informatics standards. Symbolically
published on the 250th anniversary of the officially recognized date of publication of Systema Naturae (Lin-
naeus 1758), it is our hope that this article will help demonstrate the value of electronic informatics standards
in enhancing the work of all taxonomists throughout the next quarter-millennium and beyond.

Material and methods

Specimen collection and deposition

All specimens were collected with hand nets (sometimes with the aid of rotenone or quinaldine sulphate)
by divers using Cis-Lunar® MK-5P mixed-gas, closed-circuit rebreathers, generally following protocols out-
lined by Pyle (1996c). In all cases, specimens were collected in full accordance with local government regula-
tions, and in compliance with appropriate animal care standards. Most specimens were photographed when
fresh, usually following the technique described by Randall (1961). Before exposure to formalin, tissue sam-
ples (usually the right pelvic fin) were removed from many specimens and stored in vials containing DMSO
for later analysis. Holotypes of C. abyssus, C. brevirostris, C. circumaurea, and C. degruyi are deposited in
the fish collection of the Bernice P. Bishop Museum, Honolulu (BPBM)16; the holotype of C. earina is depos-
ited in the fish collection of Muséum National d’Histoire Naturelle, Paris (MNHN)17. Paratypes are deposited
in these two insitutions, as well as The Natural History Museum, London (BMNH)18; California Academy of
Sciences, San Francisco (CAS)19; United States National Museum of Natural History, Washington, D.C.
(USNM)20; and the Western Australian Museum, Perth (WAM)21. Tissue samples were deposited for long-term
cryogenic storage at the Pacific Center for Molecular Biodiversity (PCMB)22, located at BPBM. All of these
collections have been registered with the Biodiversity Collections Index (BCI)23.

Counts, measurements and molecular analysis

Counts of median fin rays, vertebrae, and predorsal bones were taken from digitally-scanned radiograph
images (“x-rays”)24. Measurements of standard length (SL), lengths from tip of snout to origins of dorsal, pel-
vic and anal fins, dorsal- and anal-fin bases, and dorsal- and anal-fin spine lengths of specimens not bent or
otherwise distorted were calculated to the nearest tenth (0.1) of a mm from radiograph images25, and verified
against measurements taken directly from specimens. All other counts and measurements were taken directly
from specimens using a Fowler® Ultra-Cal IV 300-mm digital caliper. Measurement values were recorded to

4 · Zootaxa 1671 © 2008 Magnolia Press                                                                                           PYLE ET AL.

the nearest hundredth (0.01) of a mm, but rounded to the nearest tenth (0.1) of a mm for analyses. Methods of
counts and measurements are mostly consistent with those described in Allen & Randall (2004).

Unless otherwise indicated, specimen lengths are SL26, measured as the straight-line distance between the
anteriormost tip of the upper lip and the posterior edge of the hypural plate; head length27 is between anterior-
most tip of upper lip and posterior edge of opercle flap; body depth28 is maximum vertical distance between
belly and base of dorsal spines; body width29 is maximum width posterior to opercular opening; snout length30
is straight-line distance from anteriormost tip of upper lip to closest point on anterior fleshy edge of orbit;
orbit diameter31 is horizontal fleshy diameter; interorbital width32 is least bony width between eyes; upper jaw
length33 is straight-line distance from anteriormost tip of upper lip to ventroposteriormost tip of maxilla; cau-
dal peduncle depth34 is least depth; caudal peduncle length is horizontal distance between verticals at posterior
base of anal fin and posterior edge of hypural plate35; fin spine36 and ray37 lengths are straight-line distances
between extreme base and distal tip (including filamentous extension, if any); caudal fin length38 is horizontal
distance from posterior edge of hypural plate to a vertical at distal tip of longest ray (including filamentous
extension, if any); caudal concavity39 is horizontal distance between tips of the shortest and longest rays; pec-
toral-fin length40 is length of longest ray; pelvic fin length41 is length of longest ray; pectoral-fin ray count42
includes small splint-like, uppermost rudimentary ray; lateral-line scale count43 includes only those scales with
tubes; a separate count is provided for pored or pitted scales occurring midlaterally on caudal peduncle44;
scale-row counts above45 and below46 lateral line to origins of dorsal and anal fins (respectively) with values
that include “.5” refer to small truncate scales (if any) at bases of respective fins; gill-raker counts47 include all
rudiments and are provided as separate values for upper and lower limbs of first gill arch; last fin ray of dorsal
and anal fins sometimes branched at or near extreme base and counted as a single ray, but only when con-
nected at or near the extreme base48 (otherwise counted as two49).

Unless otherwise indicated, measurements provided in the text represent proportion of SL. Measurements
provided in the tables represent percent of SL. Counts, measurements and proportions appearing in parenthe-
ses represent ranges for paratypes, if different from the holotype. Counts represented as two values separated
by a pipe “|” represent values taken from left|right sides of specimens.

Total genomic DNA was extracted from available tissue samples using the Qiagen Dneasy® tissue kit fol-
lowing the manufacturer’s protocol. In compliance with the Barcode of Life initiative, a fragment of the cyto-
chrome oxidase I (COI) gene from the mitochondrial DNA (mtDNA) was amplified for holotypes (and some
paratypes) of all species. Primers used for amplification and sequencing were BOL-F1 (5' TCA ACY AAT
CAY AAA GAT ATY GGC AC 3') and BOL-R1 (5' ACT TCY GGG TGR CCR AAR AAT CA 3'), modified
from Wa r d et al. (2005). Each PCR had a total volume of 25 µl, containing between 10–20 ng of purified
DNA, 2.5 µl of 10x reaction buffer, 1.5 µl of 8 mM pre-mixed dNTPs, 2.5 mM of MgCl2, 0.25 µM of each
primer and two units of Taq DNA polymerase (Promega, Madison, WI). Cycling parameters for the mtDNA
were as follows: initial denaturation at 94°C for 2 min; 35 cycles of 94°C for 45 sec, 50°C for 45 sec, 72°C for
55 sec; and a final extension at 72°C for 2.5 min. The resulting reaction was purified with exonuclease I and
shrimp alkaline phosphatase enzymes. Excess oligonucleotide primers were removed through simultaneous
incubation of PCR product with exonuclease I and shrimp alkaline phosphatase (USB Corp., Cleveland OH).

Because of a broader ongoing effort by Luiz Rocha and colleagues to analyze phylogenetic patterns of
species within the genus Chromis, no attempt is made to infer evolutionary affinities of these five new species,
other than qualitative discussions about similarities with other species, if relevant.

Electronic content and hyperlinks

The online edition of this article includes extensive use of “hyperlinks” (embedded links that allow simple
redirection to online documents and resources via the internet). When viewing this online edition using hyper-
link-enabled software, clicking on the hyperlink text (dark blue in color) using the computer mouse pointer
will redirect the reader to the hyperlinked location or resource. There are two general categories of such

FIVE NEW SPECIES OF CHROMIS                                                          Zootaxa 1671 © 2008 Magnolia Press · 5

hyperlinks: internal, and external. Internal hyperlinks are used for linking between taxon names included
within this text and the full treatment (or first mention) of those taxa elsewhere within this document. Simi-
larly, internal hyperlinks are used for linking between literature citations in the text and the corresponding full
bibliographic citations in the “References” section of this aritcle. External hyperlinks are used for linking por-
tions of this document to electronic resources and information, such as online registration records of taxon
names, taxon usage instances, natural history specimen collections, literature, data files, and images. All such
external hyperlinks are denoted by a superscript number at their first use, corresponding to a Universal
Resource Locator (URL) as listed in the “Embedded External Hyperlinks” Appendix at the end of this article.
These URLs are included primarily to allow access to the online resources from the print edition, when the
online edition is not available.

Whenever possible, information made accessible through external hyperlinks conforms to current stan-
dards and protocols as developed and maintained through Biodiversity Information Standards (formerly Taxo-
nomic Databases Working Group; TDWG50). Most external hyperlinks are represented as a Life Science
Identifier (LSID51) enclosed within an HTTP proxy (to allow viewing through standard web browsers), but
others represent direct HTTP URLs or other standard protocols (e.g., “mailto”). All five new species names
established herein have been prospectively registered in ZooBank (Polaszek et al. 2005a; 2005b), the official
online registration system for the ICZN, which was launched the same day this article was published. All
other scientific names mentioned in this article have been retrospectively registered in ZooBank, and hyper-
links are directed to Taxon-Name Usage records, most of which represent taxonomic concepts corresponding
to those names as asserted herein. An Appendix at the end of this article lists most of the URLs, which are
included for the benefit of readers of the print edition. Excluded from the Appendix are all internal hyperlinks,
the three “mailto” hyperlinks associated with the email addresses of the authors, the ZooBank publication
LSID for this article that appears on the bottom of the first page, and the embedded hyperlinks associated with
the five ZooBank LSIDs for the new species described herein (all six ZooBank LSIDs can be viewed through
any standard web browser by appending the LSID to the prefix “http://zoobank.org/”).

DNA Barcodes have been deposited in GenBank52, in compliance with the Barcode of Life Data Systems
(BOLD)53 and associated Fish Barcode of Life Initiative54, and are accessible through hyperlinks associated
with PCMB numbers for each specimen. Images included in this article, as well as other relevant images, have
been made available through Morphbank55. Holotypes of C. abyssus and C. degruyi were scanned at the Uni-
versity of Texas High-Resolution X-ray CT Facility, and visualizations of those scans were provided by the
Digital Morphology (DigiMorph)56 project. References cited in this document are registered with ZooBank6,
and are available as full-page images through the Biodiversity Heritage Library (BHL)57 when copyright
allows. Descriptive data concerning type specimens of the five new species are available through the TDWG-
compliant Structure of Descriptive Data (SDD) standard. The content of this article is available as marked-up
text using the TaxonX58 and taXMLit standards. A more detailed description of how these various standards
were implemented in the generation of this document are included in a separate article, currently in prepara-
tion and planned for publication in 2008.

Chromis abyssus, new species

urn:lsid:zoobank.org:act:8BDC0735-FEA4-4298-83FA-D04F67C3FBEC

Deep Blue Chromis

(Figs. 1a –1c; Table 1; Morphbank59; DigiMorph60; GenBank61; Barcode62)

Holotype. BPBM 4086163 (81.6 mm SL), Belau (Palau) Islands; off Ngemelis Island; below and slightly N of
Blue Holes caverns (7°8'16.49"N, 134°13'18.5"E): above large rock outcrop, 110 m, hand net, R.L. Pyle, 27
April 2007 [PCMB 311364].

6 · Zootaxa 1671 © 2008 Magnolia Press                                                                                           PYLE ET AL.

TABLE 1. Proportional measurements (%SL) and counts of Chromis abyssus, new species. Values separated by a pipe
“|” are left|right or upper|lower.

Holotype                                                                Paratypes

BPBM BMNH BPBM BPBM BPBM CAS MNHN USNM USNM W A M
40861 2007.10.31.1 40855 40855 40855 225755 2007-1922 391136 391136 P.32898

FIVE NEW SPECIES OF CHROMIS                                                          Zootaxa 1671 © 2008 Magnolia Press · 7

Paratypes. BMNH 2007.10.31.165 (50.2 mm SL) [PCMB 310366]. BPBM 4085567 (3; 37.6–98.0 mm SL)
[PCMB 310068, 310269]. CAS 22575570 (64.1 mm SL) [PCMB 310571]. MNHN 2007.192272 (63.7 mm SL)
[PCMB 310473]. USNM 39113674 (2; 44.4–90.2 mm SL) [PCMB 310175]. WAM P.32898-00176 (64.5 mm SL)
[PCMB 310677]. All from same locality as holotype: sand and rubble slope with scattered rock outcroppings,
107–116 m, quinaldine and hand net, R.L. Pyle and B.D. Greene, 25 April 2007.

Diagnosis. Dorsal rays XIV,12–13 (usually 13); anal rays II,12–14 (usually 13); pectoral rays 18–19 (usu-
ally 19); spiniform caudal rays 3; tubed lateral-line scales 14–16; gill rakers 6–7+17–18 (usually 7+18; total
24–25, usually 25); body depth 1.58–1.83 in SL; color when fresh dark gray with a large iridescent dark blue
spot at center of each scale; membranes on median fins and pelvic fins opaque charcoal gray, with an irides-
cent dark blue margin on the spinous portion of the dorsal and anal fins; caudal fin mottled iridescent dark
blue and black; pectoral fins with a black ovoid spot covering the basal portion and pectoral-fin axil.

Description. Dorsal rays XIV,13 (two paratypes with XIV,12); anal rays II,13 (II,12–14); all dorsal and
anal rays branched, the last to base in some specimens; pectoral rays 19 (one paratype with 18), the upper 2
and lowermost unbranched; pelvic rays I,5; principal caudal rays 8+7=15; upper and lower procurrent caudal
rays 5, the anterior 3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 16|14
(14–16 except for one paratype with 11|15); posterior midlateral scales with a pore or deep pit 7|6 (5–8);
scales above dorsal fin to origin of dorsal fin 3.5 (3–3.5); scales below lateral line to origin of anal fin 9
(9–10); gill rakers 7+18=25 (6–7+17–18=24–25), 6 gill rakers on upper limb of gill arch in one paratype, 17
gill rakers on lower limb of gill arch in one paratype); surpaneural (predorsal) bones 3; vertebrae 12+13.

Body moderately deep, depth 1.73 (1.58–1.83) in SL, and compressed, the width 3.23 (2.65–3.56) in body
depth; head length 3.01 (2.79–3.05) in SL; dorsal profile of head with slight convexity anterior to eye, slight
concavity dorsal to eye, and slight convexity on nape; snout shorter than orbit diameter, its length 4.11
(4.00–5.74) in head length; orbit diameter 2.66 (2.28–2.90) in head length; interorbital space convex, its width
2.79 (2.59–2.92) in head length; caudal-peduncle depth 2.17 (2.15–2.59) in head; caudal-peduncle length 3.01
(3.34–4.39) in head.

Mouth terminal, small, oblique, the upper jaw forming an angle of about 40º to horizontal axis of head and
body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw
length 3.57 (3.39–4.11) in head; an outer row of conical teeth in each jaw, largest anteriorly; about 27 upper
and about 20 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer row, in 2–3
irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with rounded tip; gill
rakers long and slender, the longest on lower limb near angle about four-fifths length of longest gill filaments;
nostril with a fleshy rim, more elevated on posterior edge and located at level of middle of pupil, slightly less
than one-third distance from front of snout to base of upper lip.

Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin
of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with
free lower margin extending nearly to a vertical at posterior edge of pupil.

Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 11th
and 12th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout
containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of
middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each
membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching
about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns,
progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to
posterior margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly
from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of pelvic
spine about one-half length of spine. Origin of dorsal fin over second lateral-line scale, the pre-dorsal length
2.29 (2.24–2.54) in SL; base of spinous portion of dorsal fin contained 2.24 (2.02–2.39) in SL; base of soft

8 · Zootaxa 1671 © 2008 Magnolia Press                                                                                           PYLE ET AL.

portion of dorsal fin contained 5.75 (5.65–6.48) in SL; first dorsal spine 10.85 (7.78–11.01) in SL; second dor-
sal spine 6.64 (5.22–7.14) in SL; third dorsal spine 5.41 (4.53–5.42) in SL; fourth dorsal spine 5.00
(4.44–5.08) in SL; fifth dorsal spine 4.88 (4.39–5.05) in SL; sixth dorsal spine 4.90 (4.45–4.99) in SL; last
dorsal spine 6.19 (6.11–7.40) in SL; membranes of spinous portion of dorsal fin moderately incised; fourth
dorsal soft ray longest, sometimes with a filamentous extension, its length 4.39 (4.21–5.06) in SL; first anal
spine 10.92 (8.76–11.13) in SL; second anal spine 4.03 (3.66–4.24) in SL; first anal soft ray the longest, its
length 4.41 (4.15–4.62) in SL; caudal fin forked, without significant filamentous extensions, its length 2.89
(2.87–3.64) in SL, the caudal concavity 5.83 (4.54–8.00) in SL; fourth pectoral-fin ray longest, 2.77
(2.65–3.08) in SL; pelvic spine 5.20 (4.95–5.67) in SL; first soft ray of pelvic fin filamentous, usually reach-
ing to first through third anal-fin ray (when not broken or damaged), its length 2.86 (2.54–4.03) in SL.

Color of adults and juveniles when fresh predominantly charcoal gray, a large iridescent dark blue spot at
center of each scale (including scales on head and median fins), blue spots occupying about half of visible
area of each scale on body, decreasing in size slightly towards abdomen and ventral portion of body, blue
spots forming a near-continuous line along base of dorsal and anal fins, a vertical column of scales with irides-
cent dark blue spots extending dorsally on each interspinous membrane of dorsal fin, blue spots on scales cov-
ering soft portions of dorsal and anal fins varying in size, forming a mottled pattern of blue and black;
membranes on median fins and pelvic fins opaque charcoal gray, with an iridescent dark blue margin on
spinous portion of the dorsal fin, and a broad iridescent dark blue margin on the anal fin; caudal fin mottled
iridescent dark blue and black; pelvic-fin spine entirely iridescent dark blue, an iridescent dark blue streak on
the pelvic-fin soft rays, the filamentous extension on the pelvic fin white; pectoral fins translucent charcoal
gray with a ovoid black spot on base and axil; iris charcoal gray to black; iridescent blue fleshy orbit margin.

Color in alcohol similar to general color pattern when fresh, except charcoal gray pigment sometimes
fades to brownish gray, and iridescent dark blue is either faded to pale gray blue, or has disappeared altogether
(leaving the specimen uniform brownish gray, or sometimes charcoal gray overall).

Distribution. Only collected from the type locality; also observed at similar depths at Augulpelu Reef in
Palau. An individual of what appears to be this species was observed and photographed by Mr. Forrest Young
at 120–150 m near Manado, Sulawesi, Indonesia.

Etymology. Named abyssus, a Latinized form of the Greek noun abyssos (meaning “abyss”), to honor the
documentary film Pacific Abyss, produced by the British Broadcasting Corporation (BBC), which funded the
expedition on which the type specimens were collected. The vernacular name “Deep Blue Chromis”, a refer-
ence to both the life color of this species and the relatively (within the context of the genus) deep-dwelling
habits, is suggested instead of the more literally translated “Abyss Chromis”, so as not to imply that the spe-
cies inhabits depths commonly defined as “abyssal”.

Remarks. This species was first observed by the senior author on May 10, 1997, during a mixed-gas
rebreather dive to 120 m on the east side of Augulpelu Reef; Palau (07º 16.41' N, 134º 31.44' E). It was later
observed at the same reef at depths of 117–139 m from a submersible by Patrick L. Colin and Lori J. Bell in
February–March, 2001. In April 2005, Mr. Forrest Young and colleagues observed several individuals of this
(or a very similar) species during mixed-gas rebreather dives at depths of 120–150 m at Manado, Sulawesi,
Indonesia. The type specimens included herein are the first of this species to be collected.

From these observations, C. abyssus appears to prefer depths in excess of 115 m, staying close to the sub-
stratum among boulders and rock outcroppings, where it takes refuge in small caves and holes. Juveniles and
some subadults were also observed around limestone talus. Adults were usually observed singly or in pairs,
while subadults and juveniles were seen in small groups. All type specimens were collected in the same gen-
eral area, where the species is not uncommon. Other Chromis observed in the vicinity include three of the new
species described herein (C. brevirostris, C. degruyi, and C. earina).

Chromis abyssus is not obviously allied with any other known species of the genus. It shares some simi-
larities with a group of seven Indo-Pacific deep-dwelling Chromis species, characterized by a similar stout

FIVE NEW SPECIES OF CHROMIS                                                          Zootaxa 1671 © 2008 Magnolia Press · 9

body shape, a large eye, and usually XIV dorsal spines. In their 1985 description of C. abyssicola, Allen and
Randall noted a complex of deep-dwelling Chromis species distinguished by, among other characters, 19 or
20 pectoral rays, and 28–34 gill rakers. In addition to C. abyssicola, their complex included C. megalopsis78
Allen 1976 (now regarded as a junior synonym of C. mirationis Tanaka 1917), C. mirationis Tanaka 1917 and
C. struhsakeri Randall and Swerdloff 1973, to which we would add the subsequently named C. planesi79 Lec-
chini and Williams 2004. C. abyssus has fewer pectoral rays (18 or 19) and fewer gill rakers (24–28) than
members of this species complex, and may comprise a second grouping of deep-dwelling Chromis species,
along with C. okamuri Yamakawa and Randall 1989 from Japan, the East African C. woodsi80 Bruner and
Arnam 1979 (both easily distinguished from C. abyssus on the basis of color and certain morphological char-
acters such as number of gill rakers and tubed lateral-line scales), as well as the two new species C. circumau-
rea and C. degruyi, both described herein. Of the remaining two deep-dwelling Indo-Pacific stout-bodied
Chromis species with XIV spines, C. onumai Senou and Kudo 2007 has the high pectoral-ray count of the first
complex (19–20) and the gill-raker count of the second (25–27). C. axillaris81 (Bennett 1831) has a wide gill-
raker range (26–30) and cannot easily be placed in either complex by this character. Of the three new Chromis
with XIV dorsal-fin spines described herein (C. abyssus, C. circumaurea, and C. degruyi), each has a unique
and distinctive color pattern, and is readily distinguished from the others. Among the three, the former two (C.
abyssus and C. circumaurea) share the most similarities both in terms of morphology and in Barcode DNA
sequence data.

Chromis brevirostris, new species

urn:lsid:zoobank.org:act:2BD7CAEF-F09B-4647-B92F-62CBBC0E565C

Shortsnout Chromis

(Figs. 2a –2c; Tables 2 & 3; Morphbank82; GenBank83; Barcode84)

Holotype. BPBM 4080485 (63.7 mm SL), Caroline Islands; Puluwat Atoll; Alet Islet, S side (7°21'15.44"N,
149°10'47.03"E): outer reef drop-off with small caves and holes, 100–103 m, quinaldine and hand net, R.L.
Pyle and B.D. Greene, 11 April 2007 [PCMB 303386].

Paratypes. BMNH 2007.10.31.287 (55.2 mm SL), Belau (Palau) Islands; Augulpelu Reef, W side
(7°16'24.6"N, 134°31'26.4"E): shelf flanked by numerous small caves, 90 m, hand net, R.L. Pyle, 16 May
1997. BPBM 3767188 (55.4 mm SL), same collecting data as BMNH 2007.10.31.2, except collected on 6 May
1997. BPBM 3771389 (59.6 mm SL), same locality and depth as and BPBM 37671: cave in drop-off, rote-
none, R.L. Pyle and J.L. Earle, 12 May 1997. BPBM 3999390 (62.5 mm SL), Fiji Islands; Viti Levu Island;
outside of Suva Harbor; S of “Fish Patch”; southern wall (18°9'32.7"S, 178°23'58.44"E): sloping sand and
rubble with rock outcroppings, 87–92 m, rotenone, R.L. Pyle and D.F. Pence, 3 February 2002. BPBM
4042291 (3; 25.5–46.9 mm SL), Marshall Islands; Kwajalein Atoll, S end; Ennubuj (= Carlson) islet; ocean
side: cave within ledge, 100 m, rotenone, B.D. Greene, 30 December 2005. BPBM 4070392 (2; 50.9–59.5 mm
SL), Vanuatu; Espiritu Santo; off W coast (15°33'39.28"S, 167°16'29.82"E): steep slope with rubble and sand,
with some rocky outcrops with small caves and undercuts; many gorgonians, 60 m, quinaldine and hand net,
B.D. Greene, 7 October 2006. CAS 22575693 (2; 46.5–50.8 mm SL), same collecting data as BPBM 39993.
MNHN 2007-192394 (57.9 mm SL), Vanuatu; Espiritu Santo; off N end of Tutuba Island (15°32'28.57"S,
167°16'51.17"E): at base of outer reef drop-off ranging from 60–100 m, 100 m, rotenone and vacuum device,
R.L. Pyle, 10 October 2006. USNM 39113795 (4; 41.6–68.5 mm SL), Caroline Islands; Yap, S end; “Magic
Kingdom” (9°26'3.41"N, 138°2'5.96"E): among boulders on sloping shelf above deep drop-off, 98–100 m,
hand net, R.L. Pyle and B.D. Greene, 20 April 2007 [PCMB 307296, 307397, 307498, 307599]. WAM P.32899-
001100 (2; 58.1–61.4 mm SL), Belau (Palau) Islands; Augulpelu Reef, W side; shelf flanked by numerous small
caves (7°16'24.6"N, 134°31'26.4"E), 90 m, hand net, R.L. Pyle and J.L. Earle, 17 May 1997.

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Diagnosis. Dorsal rays XIII,13–14 (usually 14); anal rays II,15–16; pectoral rays 18–19 (usually 19);
spiniform caudal rays 2–3 (usually 3); tubed lateral-line scales 14–16; gill rakers 6–8+19–22 (usually
6–7+19–21; total 26–29); body depth 1.57–1.77 in SL; color when fresh pale lavender-tinged gray dorsally,
paler ventrally; three or four rows of scales dorsally from nape to upper caudal peduncle with gold edges;
small scales on basal sheath of dorsal fin almost entirely gold; median fin membranes lavender gray or trans-
lucent blue, suffused with gold color; iris yellow.

Description. Dorsal rays XIII,14 (two paratypes with 13); anal rays II,16 (one paratype with 15, another
paratype deformed with only 13); all dorsal and anal rays branched, the last to base; pectoral rays 18 (one
paratype with 19 on only the right side), the upper 2 and lowermost unbranched; pelvic rays I,5; principal cau-
dal rays 8+7=15 (one paratype with 7+7=14) ; upper and lower procurrent caudal rays 5, the anterior 3 (2–3)
spiniform (when 3, the anteriormost vestigial and not penetrating scales), the posterior 2 segmented and
unbranched; tubed lateral-line scales 15 (14–16); posterior midlateral scales with a pore or deep pit 7 (0–8);
scales above dorsal fin to origin of dorsal fin 4 (3.5–4); scales below lateral line to origin of anal fin 10 (9–10,
usually 10); gill rakers 8+21=29 (6–8+19–22=26–29, 6 gill rakers on upper limb of gill arch in one paratype,
22 gill rakers on lower limb of gill arch in one paratype); surpaneural (predorsal) bones 3; vertebrae 12+13.

Body moderately deep, depth 1.69 (1.57–1.77) in SL, and compressed, the width 3.04 (2.94–3.75) in body
depth; head length 3.13 (2.71–3.26) in SL; dorsal profile of head smoothly convex, sometimes with a very
slight concavity anterior to eye; snout shorter than orbit diameter, its length 4.25 (3.45–4.80) in head length;
orbit diameter 2.41 (2.00–2.66) in head length; interorbital space convex, its width 2.60 (2.56–3.39) in head
length; caudal-peduncle depth 2.05 (1.86–2.24) in head; caudal-peduncle length 3.49 (2.82–4.49) in head.

Mouth terminal, small, oblique, the upper jaw forming an angle of about 42º to horizontal axis of head and
body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw
length 3.28 (2.91–3.57) in head; teeth multi-serial, an outer row of conical teeth in each jaw, largest anteriorly;
about 25 upper and about 21 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer
row, in 2–3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with
rounded tip; gill rakers long and slender, the longest on lower limb near angle about two-thirds length of lon-
gest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle of
pupil, slightly less than one-sixth distance from front of snout to base of upper lip.

Opercle ending posteriorly in a flat spine, the tip broadly obtuse and obscured by a large scale; margin of
preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with
free lower margin extending nearly to a vertical at posterior edge of orbit.

Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 12th
and 13th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout
containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of
middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each
membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching
about four-fifths distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns,
progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to
posterior margin; small scales on basal one-sixth of pectoral fins; a median scaly process extending posteri-
orly from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of
pelvic spine slightly more than one-half length of spine.

Origin of dorsal fin over second lateral-line scale, the pre-dorsal distance 2.50 (2.37–2.76) in SL; base of
spinous portion of dorsal fin contained 2.09 (2.02–2.42) in SL; base of soft portion of dorsal fin contained
5.20 (4.69–5.38) in SL; first dorsal spine 9.41 (7.61–11.00) in SL; second dorsal spine 6.83 (5.54–8.20) in SL;
third dorsal spine 6.07 (4.51–6.98) in SL; fourth dorsal spine 5.79 (4.35–6.45) in SL; fifth dorsal spine 5.73
(4.18–6.31) in SL; sixth dorsal spine 5.67 (3.94–6.14) in SL; last dorsal spine 5.91 (4.71–6.25) in SL; mem-
branes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, usually with a fila-

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 11

mentous extension, its length 4.20 (2.55–4.34) in SL; first anal spine 9.30 (8.03–10.29) in SL; second anal
spine 4.95 (4.32–5.44) in SL; eleventh anal soft ray the longest, its length 4.99 (3.05–5.05) in SL; caudal fin
forked, its length 1.59 (1.19–2.44) in SL, the third or fourth principal caudal ray (upper and lower) with fila-
mentous extension, the caudal concavity 2.31 (1.52–4.19) in SL; fourth pectoral-fin ray longest, 2.41
(2.30–2.76) in SL; pelvic spine 4.99 (3.99–5.63) in SL; first soft ray of pelvic fin filamentous, usually reach-
ing to second anal-fin ray (when not broken or otherwise damaged), its length 3.94 (2.41–3.76) in SL.

Color of adults and juveniles when fresh pale lavender-tinged gray dorsally, fading to pale bluish-white on
thorax; pale bluish gray ventrally from anus to caudal fin; three or four rows of scales dorsally from nape to
upper caudal peduncle with gold edges; small scales on basal sheath of dorsal fin can be almost entirely gold;
faint gold tinge on posterior operculum and on scales along ventral margin anterior to pelvic-fin origin; inten-
sity of gold color variable, appearing more pearlescent underwater in some individuals; dorsal- and anal-fin
membranes pale translucent blue, suffused with gold, particularly basal 2/3 and distal portion of spinous dor-
sal fin, and basal and distal 1/3 of anal fin; soft dorsal fin gold tinged except for median portion; the extreme
distal margin of the dorsal fin, anal fin and pelvic fin pale turquoise blue; caudal fin lavender gray with faint
gold wash on membranes; pectoral fin translucent; pelvic fins white; iris yellow, fleshy membrane of orbit tur-
quoise blue, especially dorsally; interorbital space turquoise above upper lip, extending dorsally into nape in
some specimens.

Color in alcohol drab grayish light brown over most of body, darker brown above lateral line; scales above
lateral line with pale spot corresponding to gold markings in life; dorsal fin uniform brown except for a pale
gray submarginal line; anal fin brown with pale grayish brown markings and submarginal line; interorbital
region and anterior head uniform brown; thorax slightly lighter than body color, with pale ventral edge.

Distribution. This species has been observed or collected from the Marshall Islands southward to Fiji,
across the Caroline Islands from Puluwat to Palau, and south to Vanuatu. An underwater photo of what seems
to be this species (or an undescribed species very similar to C. brevirostris) taken in Bali, Indonesia, appears
as “Chromis sp.” on p. 531 of Kuiter & Debelius, 2006. C. brevirostris was not observed at Rarotonga (Cook
Islands), Kiritimati (Line Islands), or American Samoa during brief surveys of deep reefs at those localities.

Etymology. Named brevirostris, an adjective derived from the Latin words brevis (meaning “short”) and
rostrum (meaning “beak” or “snout”), in reference to the very short snout of this species relative to other spe-
cies in the genus.

Remarks. Generally abundant in its typical environment, which is characterized by steep slopes and
drop-offs at depths of about 90–120 m. Usually found in association with small holes and limestone talus,
often in aggregations ranging from a half-dozen to several dozen individuals feeding low in the water column.

Chromis brevirostris, as well as the new species C. earina (described herein) are both deep-dwelling
Chromis species with XIII dorsal spines, a deep body (1.57–1.9 in S.L.), short snout (3.45–4.8 in head), and
large eye (2.0–2.6 in head). Most counts are similar, and these two species appear to have more affinities with
each other than with any other species in the genus. They can be readily distinguished from each other on the
basis of color and by differences in general body shape (particularly the shape of the head profile, which is
smoothly convex in C. brevirostris, as opposed to slightly concave dorsal to the eye in C. earina), and dorsal-
and anal-fin soft ray counts. Chromis brevirostris has 13–14 dorsal-fin soft rays and 15–16 anal-fin soft rays
(except for one deformed individual with only 13 anal-fin soft rays); whereas C. earina has 11–12 dorsal-fin
soft rays and 12 anal-fin soft rays. Table 3 lists selected characters for four Indo-Pacific Chromis species
which seem to be most similar to C. brevirostris and C. earina. These four species, including C. alpha101 Ran-
dall 1988a, C. nigroanalis102 Randall 1988b, C. ovatiformis103 Fowler 1946, and C. pembae104 Randall and
McCosker 1992 (which we collectively and informally refer to here as the “Chromis alpha complex”) all have
XIII dorsal spines, a roughly similar body shape, and have been recorded at a depth of 40 m or greater. For
each of these species, at least one count, and all gill raker counts, range outside the respective ranges for C.
brevirostris and C. earina.

12 · Zootaxa 1671 © 2008 Magnolia Press                                                                                        PYLE ET AL.

TABLE 2. Proportional measurements (%SL) and counts of Chromis brevirostris, new species. Values separated by a
pipe “|” are left|right or upper|lower.

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TABLE 2 (continued). Proportional measurements (%SL) and counts of Chromis brevirostris, new species. Values sep-
arated by a pipe “|” are left|right or upper|lower.

14 · Zootaxa 1671 © 2008 Magnolia Press

PYLE ET AL.

TABLE 3. Selected characters of Chromis brevirostris, C. earina, and the four species comprising what is referred to
herein as the “C. alpha complex”.

Chromis circumaurea, new species

urn:lsid:zoobank.org:act:8ADC4817-8F1C-4C88-8B8A-5372A84CAEC9

Gold-rim Chromis

(Figs. 3a –3c, Ta b l e 4 ; Morphbank105; GenBank106; Barcode107)

Holotype. BPBM 40836108 (98.2 mm SL), Caroline Islands; Yap, S end; “Magic Kingdom” (9°26'3.41"N,
138°2'5.96"E): among boulders on sloping shelf above deep drop-off, 98–100 m, hand net, R.L. Pyle and B.D.
Greene, 20 April 2007 [PCMB 3080109].

Paratypes. BMNH 2007.10.31.3110 (102.4 mm SL) [PCMB 3081111]. CAS 225757112 (97.6 mm SL)
[PCMB 3078113]. MNHN 2007-1924114 (92.5 mm SL) [PCMB 3076115]. USNM 391138116 (94.2 mm SL)
[PCMB 3077117]. WAM P.32900-001118 (96.6 mm SL) [PCMB 3079119]. All with same data as holotype.

Diagnosis. Dorsal rays XIV,12–13 (usually 13); anal rays II, 13–14 (usually 13); pectoral rays 18–19;
spiniform caudal rays 3; tubed lateral-line scales 16–17; gill rakers 6–7+20–21 (total 26–27); body depth
1.68–1.86 in SL; color when fresh mahogany brown with bright yellow distally on spinous portion of dorsal
fin; soft portion of dorsal fin, caudal fin, and anal fin bright yellow.

Description. Dorsal rays XIV,13 (12 in one paratype); anal rays II,13 (14 in one paratype); all dorsal and
anal rays branched, the last to base in some specimens; pectoral rays 19 (18–19), the upper 2 and lowermost
unbranched; pelvic rays I,5; principal caudal rays 8+7=15; upper and lower procurrent caudal rays 5, the ante-
rior 3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 16|17 (16–17); posterior
midlateral scales with a pore or deep pit 8 (5–8); scales above dorsal fin to origin of dorsal fin 3.5 (3–3.5);
scales below lateral line to origin of anal fin 10 (9–10.5); gill rakers 6+21=27 (6–7+20–21= 26–27); surpaneu-
ral (predorsal) bones 3; vertebrae 12+13.

Body moderately deep, depth 1.71 (1.68–1.86) in SL, and compressed, the width 3.02 (2.93–3.22) in body
depth; head length 3.28 (3.18–3.37) in SL; dorsal profile of head with slight convexity anterior to eye, slight
concavity dorsal to eye, and slight convexity on nape; snout shorter than orbit diameter, its length 3.71
(3.82–4.30) in head length; orbit diameter 2.61 (2.35–2.63) in head length; interorbital space convex, its width
2.61 (2.54–2.74) in head length; caudal-peduncle depth 2.04 (2.02–2.04) in head; caudal-peduncle length 3.19
(2.72–3.40) in head.

Mouth terminal, small, oblique, the upper jaw forming an angle of about 40º to horizontal axis of head and
body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw
length 3.09 (2.86–3.22) in head; teeth multi-serial, an outer row of conical teeth in each jaw, largest anteriorly;
about 32 upper and about 26 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer
row, in 2–3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with
rounded tip; gill rakers long and slender, the longest on lower limb near angle about three-fourths length of
longest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle
of pupil, slightly less than one-third distance from front of snout to base of upper lip.

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 15

TABLE 4. Proportional measurements (%SL) and counts of Chromis circumaurea, new species. Values separated by a
pipe “|” are left|right or upper|lower.

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Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin
of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with
free lower margin extending nearly to a vertical at posterior edge of pupil.

Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 12th
and 13th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout
containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of
middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each
membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching
about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns,
progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to
posterior margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly
from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of pelvic
spine slightly more than one-third length of spine.

Origin of dorsal fin over fourth lateral-line scale, the pre-dorsal distance 2.31 (2.30–2.47) in SL; base of
spinous portion of dorsal fin contained 2.14 (2.09–2.18) in SL; base of soft portion of dorsal fin contained
6.16 (5.68–6.35) in SL; first dorsal spine 12.62 (9.91–10.88) in SL; second dorsal spine 6.83 (6.13–6.88) in
SL; third dorsal spine 5.52 (4.81–5.53) in SL; fourth dorsal spine 4.88 (4.73–5.09) in SL; fifth dorsal spine
4.86 (4.66–5.02) in SL; sixth dorsal spine 4.82 (4.63–5.08) in SL; last dorsal spine 6.40 (6.13–6.57) in SL;
membranes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, its length 4.20
(4.18–4.67) in SL; first anal spine 11.98 (10.50–11.05) in SL; second anal spine 3.77 (3.89–4.15) in SL; first
anal soft ray the longest, its length 4.18 (4.31–4.56) in SL; caudal fin forked, its length 3.17 (2.68–3.39) in SL,
the caudal concavity 5.46 (5.20–6.44) in SL; fourth pectoral-fin ray longest, 2.77 (2.62–2.83) in SL; pelvic
spine 5.78 (5.09–5.52) in SL; first soft ray of pelvic fin without long filamentous extension, usually not reach-
ing anal fin, its length 3.89 (3.82–4.20) in SL.

Color when fresh mahogany brown, appearing slate brown underwater; lateral line faintly brownish
cream-colored; scales below lateral line with faint brownish cream-colored broad center area, forming
approximately eight horizontal stripes visible underwater; spinous portion of dorsal fin same color as body,
becoming bright yellow distally on first spine; second through last dorsal spines and membranes abruptly yel-
low distally, yellow portion increasing from distal one-fourth of fin at third spine to distal half at eleventh
spine; soft dorsal fin entirely bright yellow except for posteriorly diminishing thin brown area basally on ante-
rior 6 rays; caudal region from posterior base of dorsal fin to posterior tip of caudal fin uniform bright yellow;
brown body color extends posterior to anal fin to lower anterior caudal peduncle; anal fin spines yellowish
white; anal fin rays and membranes bright yellow; scales along ventral margin from anus to origin anal fin
yellow; pectoral fin translucent; pelvic-fin spine translucent, medial yellow wash on anterior 3 pelvic rays,
rays otherwise translucent; pelvic-fin membranes mahogany brown basally, translucent distally; iris brown
with yellow wash.

Color in alcohol similar to fresh color, except yellow portions are much paler yellow, and brown portions
are slightly paler brown (much paler brown on thorax).

Distribution. Observed from submersibles in the Marshall Islands and Mariana Islands, but only col-
lected from Yap.

Etymology. Named circumaurea, an adjective derived from the Latin words circum (meaning “around”)
and aurea (meaning “golden, of gold”), in reference to the golden-yellow anal fin, caudal fin, and outer mar-
gin of the dorsal fin.

Remarks. This species was first observed and photographed from a submersible by Patrick L. Colin at
Enewetak in the Marshall Islands. An unconfirmed sighting and video clip of this species from the Mariana
Islands requires verification. It was observed at Yap at depths of 98–120 m, in a group of about a dozen indi-
viduals living among large (~1–2 m) rock boulders just above the upper edge of a precipitous drop-off. A

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 17

juvenile of approximately 40 mm SL was observed by the first author at a depth of 120 m, below the site
where the type specimens were collected; its color pattern was consistent with that of the adults.

Two photos appearing on p. 390 of Kuiter & Tonozuka (2001), labelled as Chromis analis120 (Cuvier
1830), bear a remarkable resemblance to C. circumaurea, but differ in number of dorsal-fin spines (XIII vs.
XIV) and color of body (paler in C. analis), caudal peduncle (dark centrally vs. entriely yellow), and central
region of caudal fin (transparent vs. yellow). This species also bears a superficial resemblance in color to
Chromis flavicauda121 (Günther 1880) from the western Atlantic Ocean, but is readily distinguished from that
species on the basis of body color (blue in C. flavicauda vs. brown in C. circumaurea), dorsal-fin rays
(XIII,11–12 vs. XIV,12–13, usually 13), anal-fin soft rays (11 vs. 13). Similarities with other deep-dwelling
species with XIV dorsal-fin spines, including the new species C. abyssus described herein, are discussed in
the Remarks section of C. abyssus.

Chromis degruyi, new species

urn:lsid:zoobank.org:act:1859B68B-340C-44F9-BEAB-D75BAED300F2

DeGruy’s Chromis

(Figs. 4a –4c; Table 5; Morphbank122; DigiMorph123; GenBank124; Barcode125)

Holotype. BPBM 40842126 (81.0 mm SL), Belau (Palau) Islands; Kayangel Atoll, W side; on outer reef drop-
off near tip of small reef extension (8°4'16.64"N, 134°40'54.52"E): rocky ledge with holes at base of steep
sandy slope with many gorgonians, 85 m, hand net, R.L. Pyle, 22 April 2007 [PCMB 3086127].

Paratypes. BMNH 2007.10.31.4128 (38.7 mm SL), Caroline Islands; Yap, S end; “Magic Kingdom”
(9°26'3.41"N, 138°2'5.96"E): deep rubble on rocky slope, 85 m, quinaldine and hand net, R.L. Pyle, 20 April
2007 [PCMB 3084129]. CAS 225758130 (38.3 mm SL), Caroline Islands; Puluwat Atoll; Alet Islet, S side
(7°21'15.44"N, 149°10'47.03"E): outer reef drop-off with small caves and holes, 100–103 m, quinaldine and
hand net, R.L. Pyle and B.D. Greene, 11 April 2007 [PCMB 3032131]. USNM 391139132 (76.6 mm SL), Belau
(Palau) Islands; off Ngemlis Island; below and slightly to the N of the Blue Holes cave system (7°8'16.49"N,
134°13'18.5"E): in coral and rubble at the base of a large boulder offset from the drop-off, 88 m, hand net,
R.L. Pyle, 27 April 2007 [PCMB 3114133]. WAM P.32901-001134 (82.4 mm SL), Belau (Palau) Islands; Ngaru-
angl Atoll, S end (8°8' 50.39"N, 134°37'3.47"E), 115 m, hand net, R.L. Pyle, 23 April 2007 [PCMB 3088135].

Diagnosis. Dorsal rays XIII–XIV,11–12 (usually XIV,12); anal rays II,11–12 (usually 12); pectoral rays
18; spiniform caudal rays 3; tubed lateral-line scales 15–17; gill rakers 7+20–21 (total 27–28); body depth
1.84–1.99 in SL; color of adults when fresh dull brownish yellow with nine thin lavender-gray stripes on side
of body, with a prominent black spot on dorsal half of pectoral-fin base.

Description. Dorsal rays XIV,12 (one paratype with XIII, another with 11); anal rays II,12 (one paratype
with 11); all dorsal and anal rays branched, the last to base in some specimens; pectoral rays 18, the upper 2
and lowermost unbranched; pelvic rays I,5; principal caudal rays 8+7=15; upper and lower procurrent caudal
rays 5, the anterior 3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 16|15
(15–17, one paratype with 17); posterior midlateral scales with a pore or deep pit 8|9 (5–9); scales above dor-
sal fin to origin of dorsal fin 3; scales below lateral line to origin of anal fin 9 (one paratype with 8); gill rakers
7+20=27 (7+20–21=27–28); surpaneural (predorsal) bones 3; vertebrae 12+13.

Body moderately deep, depth 1.84 (1.92–1.99) in SL, and compressed, the width 2.87 (2.73–3.29) in body
depth; head length 3.10 (2.95–3.18) in SL; dorsal profile of head with slight convexity anterior to eye, very
slight concavity dorsal to eye, and very slight convexity on nape; snout shorter than orbit diameter, its length
4.05 (3.63–4.38) in head length; orbit diameter 2.77 (2.12–2.95) in head length; interorbital space convex, its
width 2.73 (2.73–3.15) in head length; caudal-peduncle depth 2.18 (2.11–2.27) in head; caudal-peduncle
length 2.83 (2.69–3.37) in head.

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TABLE 5. Proportional measurements (%SL) and counts of Chromis degruyi, new species. Values separated by a pipe
“|” are left|right or upper|lower.

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 19

Mouth terminal, small, oblique, the upper jaw forming an angle of about 37º to horizontal axis of head and
body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw
length 2.91 (3.05–3.18) in head; teeth multi-serial, an outer row of conical teeth in each jaw, largest anteriorly;
about 20 upper and about 20 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer
row, in 2–3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with
rounded tip; gill rakers long and slender, the longest on lower limb near angle about three-fourths length of
longest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle
of pupil, slightly less than one-third distance from front of snout to base of upper lip.

Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin
of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with
free lower margin extending nearly to a vertical at posterior edge of pupil.

Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 13th
and 14th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout
containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of
middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each
membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching
about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns,
progressively smaller distally; small scales on caudal fin extending slightly more than two-thirds distance to
posterior margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly
from between base of pelvic fins, its length about half that of pelvic spine; axillary scale above base of pelvic
spine about one-half length of spine.

Origin of dorsal fin over third lateral-line scale, the pre-dorsal distance 2.39 (2.30–2.41) in SL; base of
spinous portion of dorsal fin contained 2.17 (2.14–2.34) in SL; base of soft portion of dorsal fin contained
6.66 (6.54–7.17) in SL; first dorsal spine 12.33 (9.51–12.14) in SL; second dorsal spine 8.15 (6.84–7.96) in
SL; third dorsal spine 5.88 (5.53–6.39) in SL; fourth dorsal spine 5.23 (5.04–5.87) in SL; fifth dorsal spine
5.08 (5.17–5.76) in SL; sixth dorsal spine 5.07 (4.93–5.79) in SL; last dorsal spine 6.27 (6.14–7.04) in SL;
membranes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, its length 4.95
(4.54–5.04) in SL; first anal spine 11.30 (10.99–13.08) in SL; second anal spine 3.84 (3.76–4.52) in SL; first
anal soft ray the longest, its length 4.48 (4.34–5.15) in SL; caudal fin forked, its length 2.84 (2.26–3.29) in SL,
the caudal concavity 5.63 (4.37–5.46) in SL; fourth pectoral-fin ray longest, 2.99 (2.79–3.20) in SL; pelvic
spine 5.36 (5.66–6.12) in SL; first soft ray of pelvic fin filamentous, usually reaching to first or second anal-
fin ray (when not broken or otherwise damaged), its length 2.94 (2.95–4.17) in SL.

Color of adults when fresh dull brownish yellow with nine thin lavender-gray stripes, some faint, the mid-
dle 4 or 5 extending onto caudal peduncle; nape area olive-brown, lighter on thorax and ventrally to anus,
becoming yellowish white; black spot slightly smaller than orbit at upper pectoral axil; faint diffuse lavender
blotch smaller than orbit on opercle edge at level of lower orbit, not apparent underwater; olivaceous with
brown stripes and greenish olive in nape area when observed underwater; spinous portion of dorsal fin olive-
brown, distal one-fourth yellowish white; basal half of soft dorsal fin dark brown with almost black outer mar-
gin, distal half very light yellowish white to translucent on some specimens; caudal fin olive-brown, inner
rays yellowish white; anal fin spines yellowish white, rays and membranes on basal half light olive-brown
becoming distally yellowish white; black blotch smaller than orbit centered in posterior distal anal fin, more
apparent in large specimens; pectoral fin translucent; pelvic-fin spine and first ray white, successive rays and
membranes on basal half olive-brown, distal half yellowish white; iris brownish yellow; juveniles bluish gray;
a bright yellow blotch on the distal half of the soft dorsal fin, covering the second through fifth dorsal soft
rays, rays 6 to last paler than anterior part of soft dorsal fin; a bright yellow stripe from posterior base of soft
dorsal fin in a band approximately the width of 2 scales continuing dorsally to tip of outer rays of dorsal lobe
of caudal fin; lower caudal rays with a similar yellow band commencing ventrally on caudal peduncle and

20 · Zootaxa 1671 © 2008 Magnolia Press                                                                                        PYLE ET AL.

extending to distal tip of lower caudal fin rays, approximately 7 median caudal rays white; third through sev-
enth anal-fin rays and intervening membranes bright yellow on distal third, succeeding rays white.
Color in alcohol similar to fresh color, except paler brown overall.

Distribution. Observed or collected throughout the Caroline Islands, from Puluwat to Palau. A Chromis
resembling the juvenile of this species was observed in Fagatele Bay in May 2001 by the senior author.

Etymology. Named degruyi to honor Michael V. DeGruy, in recognition of the sincere enthusiasm and
determination he demonstrated while attempting to collect the first adult specimen of this species.

Remarks. The habitat of this species is similar to that of other species described herein: deep outer-reef
slopes at depths of 85–120 m, usually in the vicinity of rock outcrops with small holes and caves, and around
limestone talus. It is generally not as abundant as other species of Chromis described here, usually found in
small groups and observed feeding low in the water column.

The species appears most similar to other deep-dwelling species of Chromis described previously (see
Remarks section of C. abyssus). Juveniles superficially resemble C. opercularis136 (Günther in Playfair and
Günther 1867) in color, but are easily distinguished from that species on several morphological characters
(e.g., usually XIV dorsal-fin spines in C. degruyi, vs. XIII in C. opercularis; body depth 1.84–1.99 in SL vs.
2.1–2.3 in C. opercularis), as well as adult coloration. Some aspects of the adult coloration are similar to C.
planesi Lecchini and Williams 2004, but C. degruyi differs from that species in many other aspects of adult
coloration as well as number of pectoral-fin rays (20 in C. planesi vs. 18 in C. degruyi), dorsal-fin soft rays
(usually 13 vs. usually 12), and tubed lateral-line scales (17 vs. usually 15–16).

Chromis earina, new species

urn:lsid:zoobank.org:act:269D61C2-50B3-4A8C-BEFB-D9CFBCF91BA4

Spring Chromis

(Figs. 5a –5c; Tables 3 & 6; Morphbank137; GenBank138; Barcode139)

Holotype. MNHN 2007-1921140 (63.3 mm SL), Vanuatu; Espiritu Santo; off W coast of Tutuba Island
(15°32'39.28"S, 167°16'29.82"E): near large boulder along steep slope with rubble and sand; many gorgo-
nians, 116 m, rotenone and vacuum device, R.L. Pyle and B.D. Greene, 22 October 2006 [PCMB 3131141].

Paratypes. BMNH 2007.10.31.5142 (62.5 mm SL), Vanuatu; Espiritu Santo; off W coast of Tutuba Island
(15°32'35.23"S, 167°16'49.65"E): large rock outcrop with surrounding sand and rubble, below base of large
drop-off, 116 m, hand net, B.D. Greene, 20 October 2006. BPBM 37674143 (54.0 mm SL), Belau (Palau)
Islands; Augulpelu Reef, W side (7°16'24.6"N, 134°31'26.4"E): shelf flanked by numerous small caves, 90 m,
hand net, R.L. Pyle, 7 May 1997. BPBM 37714144 (4; 48.7–67.52), same locality as BPBM 37674: cave in
drop-off, 90 m, rotenone, R.L. Pyle and J.L. Earle, 12 May 1997. BPBM 40720145 (2; 59.7–64.4 mm SL), Van-
uatu; Espiritu Santo; off N end of Tutuba Island (15°32'28.57"S, 167°16'51.17"E): at base of outer reef drop-
off ranging from 60–100 m, 100 m, rotenone and vacuum device, R.L. Pyle, 10 October 2006. CAS 225759146
(59.6 mm SL), Fiji Islands; Viti Levu Island; Suva; outside of Suva Harbor; S end of “Fish Patch”; below cave
(18°9'36.6"S, 178°23'57.6"E): sand and rubble slope with scattered outcroppings, below base of vertical reef
drop-off, 104–110 m, rotenone, R.L. Pyle, J.L. Earle, and J. Dituri, 4 February 2002. MNHN 2007-1926147
(35.7 mm SL), same collecting data as BPBM 40720. USNM 391140148 (66.3 mm SL), Vanuatu; Espiritu
Santo; off W coast of Tutuba Island (15°32'58.78"S, 167°16'40.98"E): steep slope with rubble and sand, with
some rocky outcrops with small caves and undercuts; many gorgonians, 100 m, rotenone and vacuum device,
R.L. Pyle, 16 October 2006. WA M P. 32902-001149 (53.9 mm SL), Belau (Palau) Islands; Ngemlis Island, SE
tip; "Big Drop" (7°6' 11.89"N, 134°15'2.67"E), 85 m, hand net, R.L. Pyle, 18 May 1997.

Diagnosis. Dorsal rays XII–XIII,11–12 (usually XIII, 12); anal rays II,12; pectoral rays 17–18 (usually
18); spiniform caudal rays 3; tubed lateral-line scales 13–15 (rarely 16); gill rakers 6–8+18–21 (total 26–28,

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 21

rarely 25); body depth 1.65–1.9 in SL; color when fresh pale slate blue (bright pale green in life); a white spot
(sometimes two white spots) roughly the size of a scale mid-laterally on the body; malachite green area above
orbit and in inter-orbital space and nape; dorsal and anal fins with bright distal border of pale turquoise blue.

Description. Dorsal rays XIII,12 (one paratype with XII, another with 11); anal rays II,12; all dorsal and
anal rays branched, the last to base in some specimens; pectoral rays 18 (17–18), the upper 2 and lowermost
unbranched; pelvic rays I,5; principal caudal rays 8+7=15; upper and lower procurrent caudal rays 5, the ante-
rior 3 spiniform, the posterior 2 segmented and unbranched; tubed lateral-line scales 15 (13–16, one paratype
with 16); posterior midlateral scales with a pore or deep pit 8 (4–8); scales above lateral line to origin of dorsal
fin 3; scales below lateral line to origin of anal fin 9 (8–9); gill rakers 7+21=28 (6–8+18–21=26–28, one para-
type with 25); surpaneural (predorsal) bones 3; vertebrae 12+13.

Body moderately deep, depth 1.90 (1.65–1.89) in SL, and compressed, the width 3.29 (2.71–3.70) in body
depth; head length 2.96 (2.82–3.14) in SL; dorsal profile of head with convexity anterior to eye and concavity
dorsal to eye; snout shorter than orbit diameter, its length 3.57 (3.52–4.40) in head length; orbit diameter 2.27
(2.11–2.40) in head length; interorbital space convex, its width 2.73 (2.60–3.22) in head length; caudal-pedun-
cle depth 2.35 (2.00–2.43) in head; caudal-peduncle length 3.33 (2.79–4.26) in head.

Mouth terminal, small, oblique, the upper jaw forming an angle of about 52º to horizontal axis of head and
body; posterior edge of maxilla reaching slightly beyond a vertical at anterior edge of pupil, the upper jaw
length 3.06 (2.85–3.24) in head; teeth multi-serial, an outer row of conical teeth in each jaw, largest anteriorly;
about 30 upper and about 27 lower teeth on each side of jaw; a narrow band of villiform teeth lingual to outer
row, in 2–3 irregular rows anteriorly, narrowing to a single row on side of jaws; tongue triangular with
rounded tip; gill rakers long and slender, the longest on lower limb near angle about four-fifths length of lon-
gest gill filaments; nostril with a fleshy rim, more elevated on posterior edge and located at level of middle of
pupil, slightly less than one-third distance from front of snout to base of upper lip.

Opercle ending posteriorly in a flat spine, the tip relatively obtuse and obscured by a large scale; margin
of preopercle smooth, the posterior margin extending dorsally to level of upper edge of pupil; suborbital with
free lower margin extending nearly to a vertical at posterior edge of pupil.

Scales finely ctenoid; anterior lateral line ending beneath rear portion of spinous dorsal fin (between 12th
and 13th dorsal-fin spines); head scaled except lips, tip of snout, and a narrow zone from orbit to edge of snout
containing nostrils; a scaly sheath at base of dorsal and anal fins, about two-thirds pupil diameter at base of
middle of spinous portion of dorsal fin, progressively narrower on soft portion; a column of scales on each
membrane of dorsal fin, narrowing distally, those on spinous portion of dorsal progressively longer, reaching
about two-thirds distance to spine tips on posterior membranes; scales on anal-fin membrane in two columns,
progressively smaller distally; small scales on caudal fin extending to about one-half distance to posterior
margin; small scales on basal one-fifth of pectoral fins; a median scaly process extending posteriorly from
between base of pelvic fins, its length about one-third that of pelvic spine; axillary scale above base of pelvic
spine slightly more than one-third length of spine.

Origin of dorsal fin over third lateral-line scale, the pre-dorsal distance 2.35 (2.24–2.48) in SL; base of
spinous portion of dorsal fin contained 2.32 (2.13–2.52) in SL; base of soft portion of dorsal fin contained
6.28 (5.59–7.56) in SL; first dorsal spine 10.27 (9.20–11.88) in SL; second dorsal spine 7.11 (5.91–7.43) in
SL; third dorsal spine 5.55 (4.88–6.01) in SL; fourth dorsal spine 5.62 (4.68–5.55) in SL; fifth dorsal spine
5.50 (4.68–5.51) in SL; sixth dorsal spine 5.57 (4.68–5.68) in SL; last dorsal spine 7.05 (6.17–7.21) in SL;
membranes of spinous portion of dorsal fin moderately incised; fourth dorsal soft ray longest, its length 4.25
(3.61–4.74) in SL; first anal spine 10.85 (9.93–12.43) in SL; second anal spine 3.80 (3.76–4.23) in SL; first
anal soft ray the longest, its length 4.32 (3.69–4.87) in SL; caudal fin forked, its length 2.83 (2.45–2.99) in SL,
the caudal concavity 4.49 (3.33–5.26) in SL; fourth pectoral-fin ray longest, 2.82 (2.55–2.92) in SL; pelvic
spine 5.23 (4.62–5.85) in SL; first soft ray of pelvic fin filamentous, usually reaching to first or second anal-
fin ray (when not broken or otherwise damaged), its length 2.84 (2.47–3.91) in SL.

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TABLE 6. Proportional measurements (%SL) and counts of Chromis earina, new species. Values separated by a pipe “|”
are left|right or upper|lower.

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 23

TABLE 6 (continued). Proportional measurements (%SL) and counts of Chromis earina, new species. Values separated
by a pipe “|” are left|right or upper|lower.

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Color of adults and juveniles when fresh pale slate blue (bright pale green in life), slightly darker dorsally
and slightly lighter on thorax; a white spot roughly the size of a scale mid-laterally, approximately below the
tenth dorsal-fin spine and two scale rows below the lateral line, size and shape of spot variable, occasionally
as two small spots in vertical orientation; malachite green area above orbit and in interorbital space, extending
diffusely onto nape; bright band of pale turquoise blue below orbit extending across upper lip; spinous portion
of dorsal fin color of body, with bright distal border of pale turquoise blue; soft dorsal-fin membrane pale
blue, becoming translucent on distal half; caudal fin pale slate blue, becoming lighter and distally translucent
on inner rays; caudal fin tips dark, almost black; anal fin pale slate blue, with pale turquoise blue anterior dis-
tal border; pectoral fin translucent; yellowish spot about scale size at upper pectoral axil; pelvic fin spine and
first-ray filament pale turquoise blue, membranes translucent; iris dark gray with white inner border.

Color in alcohol olive-brown with whitish margins on median fins; white spot on side of body not visible.

Distribution. Collected or observed throughout the tropical western Pacific, from Puluwat westward to
Palau, south to Papua New Guinea, Vanuatu and Fiji. It was not observed at Rarotonga (Cook Islands), Kiriti-
mati (Line Islands), or American Samoa during brief surveys of deep reefs at those localities. A single speci-
men was also recently collected by Mark Erdmann (Conservation International) in 75 m depth at Misool
Island, Raja Ampat Islands, Indonesia (G. Allen, pers. comm.).

Etymology. Named earina, a Latinized form of the Greek adjective earinos (meaning “the color of
spring”, i.e., green), in reference to the pale green color of this species in life.

Remarks. This species inhabits the same general habitat as the other new species described herein: steep
outer reef slopes and drop-offs with rocky outcrops and small caves and holes, often in association with lime-
stone talus. It is often observed in pairs or small groups, feeding low in the water column, and is generally
abundant where it is found.

Similarities with other species are discussed in the Remarks section under the account of C. brevirostris.

Acknowledgements

The following individuals or companies provided logistic and/or financial support for field collections: the
Discovery Channel (Palau, 1997); David W. Greenfield through a grant from the University of Hawai‘i (DEB-
0102745) (Fiji, 2002); Philippe Bouchet through a grant from the Sloan Foundation (Vanuatu, 2006); and the
British Broadcasting Corporation (BBC) (Caroline Islands, 2007). Additional support for deep-diving activi-
ties was provided by the Association for Marine Exploration. Claudia R. Rocha performed the DNA sequenc-
ing of tissue samples necessary to generate the COI Barcodes (as well as other sequences). The sequencing
work was supported by a grant to Brian W. Bowen from the National Science Foundation (OCE-0453167).
We wish to thank the Commissioners of the International Commission on Zoological Nomenclature (ICZN),
particularly Dr. Miguel A. Alonso-Zarazaga, for providing very useful insights concerning etymology. Arnold
Suzumoto of the Bishop Museum fish collection provided curatorial assistance, and helped to complete por-
tions of the manuscript. Loreen O’Hara created radiographs of type specimens. Neal Evenhuis provided a
large-format scanner for digitizing radiographs. Certain informatics components of this document were sup-
ported by the Pacific Basin Information Node (PBIN) of the U.S. National Biological Information Infrastruc-
ture (NBII) through a cooperative agreement with Bishop Museum. The following individuals contributed
substantially to the process of generating online resources and/or markup documents associate with this arti-
cle: Donat Agosti, Terry Catapano, Ann Devenish, Chris Freeland, Gregor Hagedorn, Robert Hanner, Julian
Humphries, Roger Hyam, Jessie Maisano, Kevin Richards, Luiz Rocha, Katja Seltmann, Dirk Steinke, Anna
Weitzman, Robert Whitton and Zhi-Qiang Zhang. David Catania, James Maclaine, Sue Morrison, Patrice Pru-
vost, Shirleen Smith, and Jeffry T. Williams assisted with catalog numbers for type specimens and with ensur-
ing online accessibility of type-specimen data.

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 25

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sophical Transactions of the Royal Society B: Biological Sciences, 360, 1847–1857.179

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Journal of Ichthyology, 36, 299–302.180

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 27

FIGURE 1. Chromis abyssus: Color photo (a181, R. Pyle) and radiograph (b182, L. O’Hara) of BPBM 40861, Holotype,
109.8 mm TL; frame from underwater video at 120 m (c183, J. Earle).

FIGURE 2. Chromis brevirostris: Color photo (a184, R. Pyle) and radiograph (b185, L. O’Hara) of BPBM 40804, Holo-
type, 103.8 mm TL; frame from underwater video at 90 m (c186, J. Earle).

FIGURE 3. Chromis circumaurea: Color photo (a187, R. Pyle) and radiograph (b188, L. O’Hara) of BPBM 40836, Holo-
type, 129.1 mm TL; frame from underwater video at 110 m (c189, J. Earle).

FIGURE 4. Chromis degruyi: Color photo (a190, R. Pyle) and radiograph (b191, L. O’Hara) of BPBM 40842, Holotype,
109.5 mm TL; BPBM 40803, juvenile, 54.5 mm TL (c192, R. Pyle).

FIGURE 5. Chromis earina: Color photo (a193, B. Greene) and radiograph (b194, L. O’Hara) of MNHN 2007-1921, Holo-
type, 86.4 mm TL; frame from underwater video at 98 m (c195, J. Earle).

28 · Zootaxa 1671 © 2008 Magnolia Press                                                                                         PYLE ET AL.

Appendix: Embedded external hyperlinks

1.         http://www.bishopmuseum.org

2.         http://zoobank.org/urn:lsid:zoobank.org:act:B8F9F80D-5798-4342-810D-D8274164F8F1

3.         http://zoobank.org/urn:lsid:zoobank.org:act:A320D8B0-30F3-4D17-ABBB-E82C6102DA54

4.         http://zoobank.org/urn:lsid:zoobank.org:act:D462EE21-1C32-46AC-8B51-39188D18536A

5.         http://zoobank.org/urn:lsid:zoobank.org:act:B07B01A1-4172-44E2-A0E6-010D7283A9D9

6.         http://www.zoobank.org

7.         http://zoobank.org/urn:lsid:zoobank.org:act:1A66BAE9-9B37-4C73-A560-BF63D0345F04

8.         http://zoobank.org/urn:lsid:zoobank.org:act:8F966156-68DE-4147-953D-74E1B3724F5F

9.         http://zoobank.org/urn:lsid:zoobank.org:act:BF53AD75-1D9E-4225-AAA3-FC39FA4FE679

10.       http://zoobank.org/urn:lsid:zoobank.org:act:D95AE369-0217-4A3D-B0E0-D6CBD6921CC6

11.       http://zoobank.org/urn:lsid:zoobank.org:act:9590A5FC-ED74-4A7A-9E59-6DB7A1638F7D

12.       http://zoobank.org/urn:lsid:zoobank.org:act:6D9436BE-BF39-4813-A11C-91599B7F580B

13.       http://zoobank.org/urn:lsid:zoobank.org:act:8AE2FCE2-A076-49F0-AFFD-6763F43BCF4F

14.       http://www.iczn.org

15.       http://www.iczn.org/iczn/

16.       http://biocol.org/urn:lsid:biocol.org:col:1001

17.       http://biocol.org/urn:lsid:biocol.org:col:1005

18.       http://biocol.org/urn:lsid:biocol.org:col:1004

19.       http://biocol.org/urn:lsid:biocol.org:col:1003

20.       http://biocol.org/urn:lsid:biocol.org:col:1002

21.       http://biocol.org/urn:lsid:biocol.org:col:1006

22.       http://biocol.org/urn:lsid:biocol.org:col:1007

23.       http://www.biodiversitycollectionsindex.org

24.       http://www.morphbank.net/?id=196937

25.       http://www.morphbank.net/?id=196938

26.       http://www.morphbank.net/?id=196939

27.       http://www.morphbank.net/?id=196940

28.       http://www.morphbank.net/?id=196941

29.       http://www.morphbank.net/?id=196942

30.       http://www.morphbank.net/?id=196943

31.       http://www.morphbank.net/?id=196944

32.       http://www.morphbank.net/?id=196945

33.       http://www.morphbank.net/?id=196946

34.       http://www.morphbank.net/?id=196947

35.       http://www.morphbank.net/?id=196948

36.       http://www.morphbank.net/?id=196949

37.       http://www.morphbank.net/?id=196950

38.       http://www.morphbank.net/?id=196951

39.       http://www.morphbank.net/?id=196952

40.       http://www.morphbank.net/?id=196953

41.       http://www.morphbank.net/?id=196954

42.       http://www.morphbank.net/?id=196955

43.       http://www.morphbank.net/?id=196956

44.       http://www.morphbank.net/?id=196957

45.       http://www.morphbank.net/?id=196958

46.       http://www.morphbank.net/?id=196959

47.       http://www.morphbank.net/?id=196960

48.       http://www.morphbank.net/?id=196961

49.       http://www.morphbank.net/?id=196962

50.       http://www.tdwg.org

51.       http://lsids.sourceforge.net

52.       http://www.ncbi.nlm.nih.gov/Genbank/

53.       http://www.barcodinglife.org

54.       http://www.fishbol.org

55.       http://www.morphbank.net

56.       http://www.digimorph.org

57.       http://www.biodiversitylibrary.org

58.       http://hdl.handle.net/10199/15417

59.       http://www.morphbank.net/Show/?id=197036

60.       http://digimorph.org/specimens/Chromis_abyssus

61.       http://www.ncbi.nlm.nih.gov/sites/entrez?term=Chromis%20abyssus&cmd=Search&db=nuccore

62.       http://www.barcodinglife.org/views/taxbrowser.php?taxon=Chromis+abyssus

63.       http://zoobank.org/urn:lsid:zoobank.org:specimen:FDE70A5C-59C3-407B-B9A6-5A9A2DA14BD1

FIVE NEW SPECIES OF CHROMIS                                                        Zootaxa 1671 © 2008 Magnolia Press · 29

64.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:E11EA6B9-5F58-4ACF-B1FF-9B7030EB4380

65.       http://zoobank.org/urn:lsid:zoobank.org:specimen:D6B9366F-B00C-4205-A6E8-933950643B94

66.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:5C149E80-12CC-40EE-83A8-4CD4D73C1C98

67.       http://zoobank.org/urn:lsid:zoobank.org:specimen:9CA82D49-9408-4A2F-99F1-AC1E8B98DAB3

68.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:0523BF75-76A9-4FEB-AF7D-5AC82F5B1BB6

69.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:AFCF1305-DB40-45B3-A83C-38525DBB2E69

70.       http://zoobank.org/urn:lsid:zoobank.org:specimen:44EA2339-5581-4A47-85D8-1AB021B34160

71.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:E393CD7D-0887-452D-9746-F4EDBE94814D

72.       http://zoobank.org/urn:lsid:zoobank.org:specimen:CF9585AB-A31E-46CB-8353-A869CA859D51

73.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:BD6CABC4-D188-4BE5-946D-F2C0E78DE52F

74.       http://zoobank.org/urn:lsid:zoobank.org:specimen:0F8CFC46-D652-49E4-BF48-9C291A6744D8

75.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:32BC8772-8D2B-40BB-8C87-AE99FB3201B3

76.       http://zoobank.org/urn:lsid:zoobank.org:specimen:B6106680-9D21-48BC-8D16-511C5CDA611E

77.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:2616392A-461D-4388-BFEC-884511C5675A

78.       http://zoobank.org/urn:lsid:zoobank.org:act:96FAD906-5A71-4127-8AF0-0FEECEBFC8AD

79.       http://zoobank.org/urn:lsid:zoobank.org:act:FCD26A8B-B378-4F22-9F61-C36EB086020A

80.       http://zoobank.org/urn:lsid:zoobank.org:act:458A31D2-1C12-4190-BCFF-5BF1638F9364

81.       http://zoobank.org/urn:lsid:zoobank.org:act:8AAE51D4-2523-439D-B470-ADC0E6ABCC3A

82.       http://www.morphbank.net/Show/?id=197037

83.       http://www.ncbi.nlm.nih.gov/sites/entrez?term=Chromis%20brevirostris&cmd=Search&db=nuccore

84.       http://www.barcodinglife.org/views/taxbrowser.php?taxon=Chromis+brevirostris

85.       http://zoobank.org/urn:lsid:zoobank.org:specimen:6C53E362-23A2-4B3A-8478-D7E6692D3D9B

86.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:5BC49AC5-9737-45DF-A321-77FED26EB1A2

87.       http://zoobank.org/urn:lsid:zoobank.org:specimen:567A113F-99E4-450D-AFED-2716675D3A58

88.       http://zoobank.org/urn:lsid:zoobank.org:specimen:035F19C1-82D4-420D-91AD-FEFDDB68E428

89.       http://zoobank.org/urn:lsid:zoobank.org:specimen:B8FADC07-3BFE-46D1-8A6E-219E0B7AB71F

90.       http://zoobank.org/urn:lsid:zoobank.org:specimen:A9616045-3D20-470D-9264-1C27F8B422AD

91.       http://zoobank.org/urn:lsid:zoobank.org:specimen:9B55732E-1E5C-4D84-B106-723907996F6F

92.       http://zoobank.org/urn:lsid:zoobank.org:specimen:CE331ED9-3B61-40A4-B6E5-74CE559D887B

93.       http://zoobank.org/urn:lsid:zoobank.org:specimen:3D032426-87BE-42BE-A6A3-6048FB19324C

94.       http://zoobank.org/urn:lsid:zoobank.org:specimen:66822D0E-8BB7-424A-B1C3-F79FD70B0CB0

95.       http://zoobank.org/urn:lsid:zoobank.org:specimen:EFEC8CDD-C3D3-4178-8CAF-6B29AE21C053

96.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:4756AFA4-300B-4703-9B38-8B9C12351459

97.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:666613F1-CAAD-4F05-991B-49D314B98693

98.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:515F230E-8337-4077-B5BC-C640C017304B

99.       http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:AF0BD824-9B9C-4FDC-857A-2F9C00B5F7E8

100.     http://zoobank.org/urn:lsid:zoobank.org:specimen:80265A9B-5B39-43D6-B85C-9ED08F552A2F

101.     http://zoobank.org/urn:lsid:zoobank.org:act:6D0B1F3B-D066-4CD7-AF88-69863CB5A50E

102.     http://zoobank.org/urn:lsid:zoobank.org:act:D29D5552-2629-48B1-A9F3-E965CE6A978A

103.     http://zoobank.org/urn:lsid:zoobank.org:act:D12C58D1-0E72-47EE-B005-1556D07B6449

104.     http://zoobank.org/urn:lsid:zoobank.org:act:B96D923C-AA58-43BF-ADCE-6B1B28E4135A

105.     http://www.morphbank.net/Show/?id=197038

106.     http://www.ncbi.nlm.nih.gov/sites/entrez?term=Chromis%20circumaurea&cmd=Search&db=nuccore

107.     http://www.barcodinglife.org/views/taxbrowser.php?taxon=Chromis+circumaurea

108.     http://zoobank.org/urn:lsid:zoobank.org:specimen:AC204B49-93B7-4BEE-890B-7BF07C1EF592

109.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:F8DCA388-8031-4856-ACB0-CE7D1149FCC1

110.     http://zoobank.org/urn:lsid:zoobank.org:specimen:03C05B9B-816B-4904-AF54-AF55FF33CA83

111.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:07410D4D-50AD-4300-9178-C54FA8ECC1FF

112.     http://zoobank.org/urn:lsid:zoobank.org:specimen:7990C0F7-32A4-4BDB-8E6E-ED6B8B093039

113.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:5082B666-AA22-46B0-8D2D-93FEEF605054

114.     http://zoobank.org/urn:lsid:zoobank.org:specimen:E0409C2B-32A1-48E7-A8B4-DC3BAC151577

115.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:22E8BFEA-8659-4649-899A-1136F494EE3C

116.     http://zoobank.org/urn:lsid:zoobank.org:specimen:1ED95527-8AE3-495E-8564-8B72D8931A8D

117.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:2CE51FBF-12EB-4160-8625-07BC27318C2C

118.     http://zoobank.org/urn:lsid:zoobank.org:specimen:2324B43A-74A1-4AB3-A9B2-D38D7227BC10

119.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:22FB7FBA-56BF-42EB-BD08-56E702E805DF

120.     http://zoobank.org/urn:lsid:zoobank.org:act:EED194A3-93DC-47EA-A3A4-D6DCFFBAAAFA

121.     http://zoobank.org/urn:lsid:zoobank.org:act:80F113ED-4499-49BB-A07D-91C8142EE830

122.     http://www.morphbank.net/Show/?id=197039

123.     http://digimorph.org/specimens/Chromis_degruyi

124.     http://www.ncbi.nlm.nih.gov/sites/entrez?term=Chromis%20degruyi&cmd=Search&db=nuccore

125.     http://www.barcodinglife.org/views/taxbrowser.php?taxon=Chromis+degruyi

126.     http://zoobank.org/urn:lsid:zoobank.org:specimen:6A1A460A-6616-4B52-91E7-327A65E48BB5

127.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:D4B3F792-06D1-4065-BBDE-9B0FC6EC0D54

128.     http://zoobank.org/urn:lsid:zoobank.org:specimen:548DCE90-1F29-4FC3-974D-C846794696C3

129.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:5EC1BA9E-E6FF-4724-B7A3-23A296A3879C

30 · Zootaxa 1671 © 2008 Magnolia Press                                                                                        PYLE ET AL.

130.     http://zoobank.org/urn:lsid:zoobank.org:specimen:C38EB0CD-5966-46E7-B42A-20D8E4B46813

131.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:06BDFA7E-6051-4F17-A330-77B567425684

132.     http://zoobank.org/urn:lsid:zoobank.org:specimen:1C5D5286-0911-49E9-84F3-4AE5D879236A

133.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:1CC793B5-A098-422F-9E54-C76BA9EB0FB7

134.     http://zoobank.org/urn:lsid:zoobank.org:specimen:30387811-C0AF-456C-99C7-838F7945F0F0

135.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:CC570EE5-B7A6-4E0C-96E5-5A34FB6D4BFF

136.     http://zoobank.org/urn:lsid:zoobank.org:act:4B837435-740D-471D-AC13-F08F60BC7429

137.     http://www.morphbank.net/Show/?id=197040

138.     http://www.ncbi.nlm.nih.gov/sites/entrez?term=Chromis%20earina&cmd=Search&db=nuccore

139.     http://www.barcodinglife.org/views/taxbrowser.php?taxon=Chromis+earina

140.     http://zoobank.org/urn:lsid:zoobank.org:specimen:DF0CB620-B9AF-4291-8EB1-368E3D1D4D92

141.     http://nsdb.bishopmuseum.org/urn:lsid:bishopmuseum.org:bioobject:8E7BE7B7-2625-4107-A36A-DCCC13A9AEAD

142.     http://zoobank.org/urn:lsid:zoobank.org:specimen:96E8ED61-661D-45D4-9C94-77CE2EC77BC3

143.     http://zoobank.org/urn:lsid:zoobank.org:specimen:47D7B9D1-2562-4095-96EB-214EE3EBBAF6

144.     http://zoobank.org/urn:lsid:zoobank.org:specimen:773E9624-03EF-4D71-BEE7-A3D0A508ACD1

145.     http://zoobank.org/urn:lsid:zoobank.org:specimen:A4FE9A62-D16C-4545-8FE6-CADAAC8DB847

146.     http://zoobank.org/urn:lsid:zoobank.org:specimen:9BEECF03-7EB8-4FDD-8B4E-37C729242F3B

147.     http://zoobank.org/urn:lsid:zoobank.org:specimen:106E34F7-3CD7-4107-A386-AB5AF88C55D3

148.     http://zoobank.org/urn:lsid:zoobank.org:specimen:E55D3CF5-1712-440E-8B53-6F12E623DD83

149.     http://zoobank.org/urn:lsid:zoobank.org:specimen:9510FEC7-2215-4DAB-BE04-53D8560AA041

150.     http://zoobank.org/urn:lsid:zoobank.org:pub:550BC54C-E9B8-41AB-AA10-C2CA662910E4

151.     http://zoobank.org/urn:lsid:zoobank.org:pub:2549669A-B51B-43E4-9913-EA049A9A44F2

152.     http://zoobank.org/urn:lsid:zoobank.org:pub:342F642C-3FFD-4E8F-AB01-C04EC901F377

153.     http://zoobank.org/urn:lsid:zoobank.org:pub:707B0374-DE35-45D7-8A32-42B46E28A51B

154.     http://zoobank.org/urn:lsid:zoobank.org:pub:77E4A19F-3684-46CD-969F-0E1C561297ED

155.     http://zoobank.org/urn:lsid:zoobank.org:pub:56207A32-8C72-4E4E-96D2-0AE7A3BBFCFE

156.     http://zoobank.org/urn:lsid:zoobank.org:pub:5610B9B8-90CD-49D5-9977-185D8812BC5C

157.     http://zoobank.org/urn:lsid:zoobank.org:pub:145951D6-6701-4284-8A9D-E43EBF99B9AC

158.     http://zoobank.org/urn:lsid:zoobank.org:pub:262A0AB9-5FCB-454F-B539-8658D00C6A33

159.     http://zoobank.org/urn:lsid:zoobank.org:pub:298E231C-D043-45EF-B58B-D671F5E99899

160.     http://zoobank.org/urn:lsid:zoobank.org:pub:2EC2EBB9-D3F1-430B-9DD9-A704A09BF72D

161.     http://zoobank.org/urn:lsid:zoobank.org:pub:452E1D3D-84DA-4E56-B17B-C2491A977F59

162.     http://zoobank.org/urn:lsid:zoobank.org:pub:DAF92858-1F39-46F3-9B96-EF326179EB2A

163.     http://zoobank.org/urn:lsid:zoobank.org:pub:2C6327E1-5560-4DB4-B9CA-76A0FA03D975

164.     http://zoobank.org/urn:lsid:zoobank.org:pub:FE3E9DC8-8D56-4095-8A01-E0B8BEE2178E

165.     http://zoobank.org/urn:lsid:zoobank.org:pub:3F97F9F1-3156-4D30-AD7C-1F5BF51DD609

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184.     http://www.morphbank.net/?id=196553

185.     http://www.morphbank.net/?id=196554

186.     http://www.morphbank.net/?id=196964

187.     http://www.morphbank.net/?id=196619

188.     http://www.morphbank.net/?id=196616

189.     http://www.morphbank.net/?id=196965

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191.     http://www.morphbank.net/?id=196657

192.     http://www.morphbank.net/?id=196666

193.     http://www.morphbank.net/?id=196681

194.     http://www.morphbank.net/?id=196680

195.     http://www.morphbank.net/?id=196966

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