Review of Australian Species of Paracheilinus Fourmanoir (Teleostei: Labridae), with Description of a New Species from the Great Barrier Reef and Coral Sea

Abstract

Australian species of the cirrhilabrin labrid genus Paracheilinus are reviewed. Four species of Paracheilinus are report- ed from Australian waters: P. amanda, new species, from Flora, Holmes, and Osprey Reefs, Coral Sea, off northeast Queensland, and Harrier Reef, Great Barrier Reef; P. filamentosus from Lizard Island, Great Barrier Reef; P. flavianalis from Evans and Flinders Shoals, Timor Sea, off northeast Darwin, Northern Territory, and Ashmore, Scott, Seringapatam, and Hibernia Reefs in the north-western shelf of Western Australia; and P. nursalim from Flinders Shoal, Timor Sea, off northern Darwin, Northern Territory. Paracheilinus amanda, new species, has previously been confused for P. rubricaudalis from Melanesia, but molecular analysis of mitochondrial COI recovers both species as re- ciprocally monophyletic lineages, differing from each other by 1–1.2% in genetic distance. They further differ in as- pects of live coloration of terminal phase (TP) males. Both species are allopatric and do not overlap in distribution. The new species is described on the basis of six specimens: the holotype and two paratypes from Harrier Reef, Great Barrier Reef, one paratype from Flora Reef, Coral Sea, and from two paratypes collected off Hula in southern Papua New Guinea, along the north-western margin of the Coral Sea. The discovery of P. nursalim in Australia represents a new and significant range extension from previous locality records of West Papua and Ambon Bay. Paracheilinus is rediagnosed, and keys, diagnoses, photographs, and Australian distribution records are presented for all species herein.

Full text

Review of Australian Species of Paracheilinus Fourmanoir (Teleostei:
Labridae), with Description of a New Species from the Great Barrier Reef
and Coral Sea
Yi-Kai Tea
1,2
and Fenton Walsh
3
Australian species of the cirrhilabrin labrid genus Paracheilinus are reviewed. Four species of Paracheilinus are report-
ed from Australian waters: P. amanda, new species, from Flora, Holmes, and Osprey Reefs, Coral Sea, off northeast
Queensland, and Harrier Reef, Great Barrier Reef; P. filamentosus from Lizard Island, Great Barrier Reef; P. flavianalis
from Evans and Flinders Shoals, Timor Sea, off northeast Darwin, Northern Territory, and Ashmore, Scott,
Seringapatam, and Hibernia Reefs in the north-western shelf of Western Australia; and P. nursalim from Flinders
Shoal, Timor Sea, off northern Darwin, Northern Territory. Paracheilinus amanda, new species, has previously been
confused for P. rubricaudalis from Melanesia, but molecular analysis of mitochondrial COI recovers both species as re-
ciprocally monophyletic lineages, differing from each other by 1–1.2% in genetic distance. They further differ in as-
pects of live coloration of terminal phase (TP) males. Both species are allopatric and do not overlap in distribution.
The new species is described on the basis of six specimens: the holotype and two paratypes from Harrier Reef, Great
Barrier Reef, one paratype from Flora Reef, Coral Sea, and from two paratypes collected off Hula in southern Papua
New Guinea, along the north-western margin of the Coral Sea. The discovery of P. nursalim in Australia represents a
new and significant range extension from previous locality records of West Papua and Ambon Bay. Paracheilinus is
rediagnosed, and keys, diagnoses, photographs, and Australian distribution records are presented for all species
herein.
THE labrid tribe Cirrhilabrini includes just over 100
species of reef-associated fishes (Parenti and Randall,
2018;Tea et al., 2021) distributed throughout the
Indo-Pacific. Five genera are currently recognized, viz.,
Cirrhilabrus (65 species; Tea et al., 2021,2022a), Paracheilinus
(20 species; Allen et al., 2016), Pseudocheilinops (1 species;
Schultz et al., 1960), Pseudocheilinus (7 species; Randall,
1999), and Pteragogus (10 species; Parenti and Randall, 2018).
The tribe also previously contained the monotypic genus
Conniella, but the genus was recently placed within the syn-
onymy of Cirrhilabrus on the basis of molecular and morpho-
logical data (Tea et al., 2021,2022b). Except for Pteragogus,
whose species are cryptically marked and well suited for cam-
ouflage in seagrass and macroalgal reefs, the cirrhilabrin lab-
rids are typically very colorful and occur within or in the
vicinity of coral reefs. Few, however, are as spectacular as
those from the genera Cirrhilabrus and Paracheilinus, the lat-
ter especially so.
Species of Paracheilinus are small (usually under 70 mm
SL), exuberantly colored fishes most known for their stun-
ning nuptial displays, in which males temporarily “flash”
iridescent colors, often complemented by rapid swimming
and fully erect median fins. This behavior is most apparent
a few hours before sunset, though males will readily display
throughout the day, especially when spurred by agonistic
encounters with conspecific and congeneric males (Allen
et al., 2016). They are protogynous sequential hermaphro-
dites, forming large aggregations comprising anywhere be-
tween tens to hundreds of individuals, with females greatly
outnumbering males. Like its sister genus Cirrhilabrus,Para-
cheilinus typically inhabit rubble slopes near the outer reef
edges of coral reefs, swimming several meters above the bot-
tom to pick off zooplankton and other tiny invertebrates.
Although species of Paracheilinus are popular subjects for
underwater photography, their preference for soft sediment
and rubble slopes means that they are often overlooked in
faunistic surveys. These habitats are often ignored by divers
(De Brauwer et al., 2017;De Brauwer and Burton, 2018), re-
sulting in the patchy distribution records for many species
of Paracheilinus, as well as failure to detect potentially new
ones.
Although Paracheilinus occurs widely throughout the
Indo-Pacific, the distribution of its species is highly asym-
metrical across this range. The highest diversity of species
occurs in the Coral Triangle, particularly along the Indo-
Australian Archipelago. In their review of the genus, Allen
et al. (2016) recognized 20 species as valid, three of which
they described as new. Thirteen of these occur in the Indo-
Australian Archipelago, viz., P. alfiani,P. angulatus,P. carpen-
teri,P. cyaneus,P. filamentosus,P. flavianalis,P. lineopunctatus,
P. nursalim,P. paineorum,P. rennyae,P. togeanensis,P. walton,
and P. xanthocirritus. Four species occur in the Western Indi-
an Ocean and the Red Sea, viz., P. attenuatus,P. hemitaenia-
tus,P. piscilineatus, and P. octotaenia.Paracheilinus mccoskeri
is widespread in the Indian Ocean, ranging from the east Af-
rican coast to the Andaman Sea, Sumatra, and Bali. The re-
maining two species are confined to oceanic islands of the
western Pacific. They are P. bellae from the Ryukyus Archi-
pelago and Micronesia, and P. rubricaudalis from Melanesia.
Despite the Indo-Australian Archipelago being the center
of diversity for this genus, the distribution and occurrence
of Paracheilinus in Australia and its remote territories are not
1
Ichthyology, Australian Museum Research Institute, 1 William Street, Sydney, New South Wales 2010, Australia; Email: yi-kai.tea@austmus.
gov.au. Send correspondence to this address.
2
School of Life and Environmental Sciences, University of Sydney, New South Wales 2006, Australia.
3
P.O. Box 389, Kuranda, Queensland 4881, Australia.
Submitted: 10 March 2023. Accepted: 27 May 2023. Associate Editor: M. T. Craig.
!2023 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/i2023019 Published online: 14 September 2023
Ichthyology & Herpetology 111, No. 3, 2023, 397–415
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well understood. Allen et al. (2016) reported P. filamentosus,
P. flavianalis, and P. rubricaudalis as extant in Australia. Ex-
cept for P. flavianalis, which was described from type materi-
al collected off the north-western shelf of Australia, the
status of the other two species in Australia has been doubt-
ful. Distribution records for P. filamentosus in Allen et al.
(2016) include north-eastern Papua New Guinea, the Solo-
mon Islands, and the northern Great Barrier Reef of Austra-
lia; however, the Australian record appears to be spurious,
with no photographs or examined material corroborating
this distribution. An extensive search of fish collections by
the first author of this study yielded only a single, previously
unexamined specimen of P. filamentosus collected within
Australian waters, from Lizard Island in the northern Great
Barrier Reef.
The status of P. rubricaudalis in Australia is similarly con-
tentious. The species is known from Australia based only on
a handful of specimens collected in the Coral Sea and the
Great Barrier Reef. These specimens, however, differ from
topotypical examples of P. rubricaudalis from Melanesia in
live coloration of terminal phase (TP) males. The Australian
specimens are compared with P. rubricaudalis from Fiji and
northern Papua New Guinea, and are herein described as a
new species, P. amanda.
Finally, examination of labrid specimens collected from
Flinders Shoal, Timor Sea, approximately 300 kilometers
north of Darwin in the Northern Territory yielded a speci-
men of Paracheilinus unassignable to any of the aforemen-
tioned species. Closer examination of color photographs
and preserved material agrees well with P. nursalim, a species
previously known only from Ambon in the Malukus and
the Bird’s Head Peninsula in West Papua.
The purpose of this study is to provide a review of Paracheili-
nus occurring in Australia, of which there are four: P. a m a n d a ,
new species, from Flora, Holmes, and Osprey Reefs, Coral Sea,
off northeast Queensland, and Harrier Reef, Great Barrier Reef;
P. f i l a m e n t o s u s ,knownfromasinglespecimencollectedfrom
Lizard Island, northern Great Barrier Reef; P. f l a v i a n a l i s from
Evans and Flinders Shoals, Timor Sea, off northern Darwin,
Northern Territory, and Ashmore, Scott, Seringapatam, and
Hibernia Reefs in the north-western shelf of Western Australia;
and P. n u r s a l i m from Flinders Shoal, Timor Sea, off northern
Darwin, Northern Territory. Paracheilinus is rediagnosed on the
basis of new character accounts, and keys, diagnoses, photo-
graphs, and Australian distribution records are presented for all
species.
MATERIALS AND METHODS
Meristics, morphometrics, coloration description, and specimen
deposition.—Methods for counting and measuring follow Ran-
dall and Allen (2003) and Allen et al. (2016),exceptgillraker
counts are presented as upper (epibranchial) þlower (cerato-
branchial); the angle raker is included in the second count.
Counts of lateral-line scales are given in two parts, the dorso-
anterior series and the midlateral peduncular series. The latter
series consists of a larger pored scale overlapping the caudal-
fin base, which we include in the count. In the description of
the new species that follows, data are presented first for the
holotype, followed by the range of minimum–maximum val-
ues of the paratypes in parentheses where different. Where
counts were recorded bilaterally, both counts are given and
separated from each other by a slash; the first count presented
is the left count. Morphometric values are expressed as per-
centage of standard length (SL; Table 1). Caudal-fin length was
measured from the base of the hypural crease to the tips of the
central rays (not including filaments where present). Caudal
concavity is the horizontal distance between verticals at the
tips of the shortest and longest caudal-fin rays. Osteological
details were taken from cleared and stained specimens, and ra-
diographs of examined specimens; for the new species, this in-
cludes the paratypes.
We follow the terminology provided by Allen et al. (2016)
in determination of the male stripe pattern of Paracheilinus,
which can be useful in diagnosing several species groups (Fig.
1). Note that these patterns do not necessarily reflect phylo-
genetic relationships within the genus. Where relevant, we
refer to horizontal or near horizontal marks, mostly on the
sides, as stripes, and curved or oblique marks on the head
and fins as bands. Unless specified, color patterns are de-
scribed from TP males.
Pattern A (sensu Allen et al., 2016;Fig. 1A1) consists of
three bands on the head, two originating from the posterior
edge of the orbit and extending to the opercle margin, and
one originating on the upper lip and passing along the
lower edge of the orbit to the lower margin of the opercle.
The body pattern consists of two primary stripes that run
along the middle portion of the side to the caudal peduncle
with several shorter, intervening secondary stripes. Allen
et al. (2016) made note of two variations in this basic pat-
tern. The first consists of scattered small dark spots and bro-
ken dashes on the side of the body in the spaces between
the primary stripes. This pattern is especially evident in fe-
males and young males, and are found in P. cyaneus,P. nursa-
lim, and P. walton. To differentiate this from pattern A, we
use the term pattern A’ (Fig. 1A2). The second variation ap-
plies only to P. alfiani, in which the two primary stripes are
somewhat expanded and diffuse on the posterior half of the
body. Since this character appears to be autapomorphic for
P. alfiani, we do not propose secondary terminology for this
pattern.
Pattern B (sensu Allen et al., 2016;Fig. 1B)consistsofa
series of anterior stripes and bands arranged in the follow-
ing manner: a short band from front of orbit to snout tip, a
band from upper lip, passing along the lower edge of orbit
to the lower opercular edge and continuing on the side of
the breast to above the pelvic fins, another upward-slant-
ing band from the dorsoposterior corner of orbit to the
opercular edge and continuing for a short distance on the
adjacent upper body, and another from midposterior edge
of orbit to middle opercular edge, and continuing on the
side of body below the pectoral-fin base to the caudal pe-
duncle. Three additional stripes are present above this
main stripe: the first along the dorsal-fin base, from nape
to upper edge of the caudal peduncle; the second and third
much shorter, situated just below, in parallel but displaced
from each other. The caudal fins of species bearing pattern
B are rounded and consist of two concentric bands follow-
ing the contours of the fin shape; the first situated on the
basal third of the fin, and the second on the outermost
edge. Allen et al. (2016) did not include the stripe along
the dorsal-fin base and caudal fin as part of this pattern,
but we here include it as an additional level of detail diag-
nosing pattern B.
The Western Indian Ocean species P. attenuatus,P. hemi-
taeniatus, and P. piscilineatus share an unusual coloration
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pattern in which the base of the pectoral fin is partially
ringed in an arc, with the posterior end extending to the
caudal peduncle in at least P. attenuatus and P. piscilineatus.
Allen et al. (2016) made note of this pattern in the latter two
species, though it also occurs in P. hemitaeniatus. While all
three are easily identified based on other unique characters
(see Allen et al., 2016), the sharing of the “shepherd’s crook”
stripe on the pectoral-fin base suggests its usefulness as a
Fig. 1. Schematic diagrams illustrating the body stripe pattern in species of Paracheilinus. (A1) Pattern A; (A2) Pattern A’. Note spot-band pattern
derived from Pattern A; (B) Pattern B. Note caudal-fin markings, which do not typically vary for species sharing Pattern B; (C) Pattern C. Note “shep-
herd’s crook” marking at pectoral-fin base. Illustration by J. W. Soong.
Table 1. Proportional measurements for type specimens of Paracheilinus amanda, new species, expressed as percentage of the standard length.
QM I.39758 AMS I.50116-001 WAM P.33973.001 ZRC 64175
Holotype Paratypes
Sex Male Male Male Male Male Female
Standard length (mm) 47.6 44.4 43.7 37.3 47.6 32.3
Body depth 31.3 26.1 29.1 28.2 27.5 28.5
Body width 15.1 13.7 12.6 14.5 14.3 12.7
Head length 32.4 30.6 31.6 33.5 33.8 35.6
Snout length 8.2 7.7 7.8 8.3 7.8 8.0
Orbit diameter 9.0 9.5 8.5 11.0 7.6 11.1
Interorbital width 9.7 9.7 9.8 11.0 9.7 8.4
Upper-jaw length 8.2 7.9 6.2 8.0 6.3 5.6
Caudal-peduncle depth 14.1 13.3 14.0 14.8 13.7 12.4
Caudal-peduncle length 18.1 18.5 14.2 14.7 16.0 16.4
Predorsal length 31.3 32.4 33.9 34.0 33.6 35.3
Preanal length 52.7 55.2 57.4 60.6 56.9 59.7
Prepelvic length 33.6 34.9 33.6 37.5 38.9 36.5
Dorsal-fin base 55.9 54.5 51.0 51.0 52.7 52.0
First dorsal spine 3.4 2.9 3.4 3.2 2.3 3.7
Longest dorsal spine (9
th
) 17.0 12.2 13.3 13.1 16.6 13.0
Longest dorsal ray (1
st
) 48.7 27.7 21.5 18.2 54.4 17.6
Last dorsal ray 9.0 9.5 8.0 6.4 13.9 10.5
Anal-fin base 28.4 26.6 28.6 27.6 27.5 23.2
First anal spine 6.3 6.8 5.0 4.3 7.4 3.7
Second anal spine 8.6 7.2 7.1 6.7 9.5 7.4
Third anal spine 10.9 9.0 9.2 9.1 10.5 8.4
First anal ray 15.8 11.0 12.4 12.3 16.6 11.5
Longest anal ray (6
th
–7
th
) 23.9 14.2 17.4 13.4 26.9 12.4
Caudal-fin length 27.5 22.3 24.9 21.7 24.7 25.1
Pectoral-fin length 20.8 20.7 22.0 21.2 20.4 21.4
Pelvic-spine length 10.5 7.9 9.6 7.8 7.8 9.6
Pelvic-fin length 16.0 11.0 14.0 12.3 14.9 13.3
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diagnostic pattern separating them from the remaining spe-
cies of Paracheilinus. We refer to this as pattern C (Fig. 1C). A
further set of species possess body patterns that do not agree
with patterns A, A’, B, or C. They are P. bellae,P. lineopuncta-
tus,P. octotaenia, and P. rennyae. These species are, however,
easily recognized based on their unique coloration details
and combination of morphological characters (see detailed
diagnoses in Allen et al., 2016).
Type specimens of the new species are deposited in the
Australian Museum, Sydney (AMS), Queensland Museum,
Brisbane (QM), the Western Australian Museum, Perth
(WAM) and the Zoological Reference Collection of the Lee
Kong Chian Natural History Museum at the National Uni-
versity of Singapore (ZRC). Comparative specimens of Para-
cheilinus were examined from material deposited at AMS,
QM, WAM, Museums Victoria, Melbourne (NMV), Museums
and Art Galleries of the Northern Territory, Darwin (NTM),
and ZRC. Institutional codes follow Sabaj (2020).
Taxon sampling, sequencing, and phylogenetic analysis.—Tissue
samples of the new species were obtained from the right pel-
vic fin of the AMS paratype (AMS I.50116-001), preserved in
100% ethanol, and stored at –208C prior to extraction. DNA
was extracted using the DNeasy Blood and Tissue kit (Qia-
gen) following the manufacturer’s protocol. Mitochondrial
cytochrome c oxidase subunit I (COI) was amplified from
extracted gDNA using the polymerase chain reaction (PCR).
Primer sets and PCR conditions follow Chang et al. (2017).
Sanger sequencing was outsourced to the Australian Ge-
nome Research Facility (Canberra, Australia). Forward and
reverse contigs were aligned and trimmed separately using
GENEIOUS Prime 2019.1.1 (Biomatters). The COI sequence
of the new species was then combined with those of other
species of Paracheilinus used in Allen et al. (2016),alignedusing
the MUSCLE v3.8.31 algorithm (Edgar, 2004), and re-analyzed
using maximum likelihood (described below). Cirrhilabrus isos-
celes was used as the outgroup. The final sequence alignment
consisted of 642 base pairs. The following newly generated se-
quences and their GenBank accession numbers not previously
in Allen et al. (2016) were used in this study: Paracheilinus
amanda,newspecies(OQ605966);P. f l a v i a n a l i s (OQ600543);
and P. r u b r i c a u d a l i s (OQ600403–06). Accession numbers for all
comparative Paracheilinus and outgroup taxa are as listed in
Allen et al. (2016).Themolecularsequencealignmentfileis
provided in the supplemental material (see Data Accessibility).
We analyzed the molecular data set using maximum like-
lihood in IQ-TREE v2.1.3 (Nguyen et al., 2015). Branch sup-
port was assessed using ultrafast bootstrapping (UFBS)
approximation with 1000 replicates (Hoang et al., 2017)
and the Shimodaira-Hasegawa-like approximate likelihood-
ratio test (SH-aLRT; Guindon et al., 2010) with 1000 repli-
cates. Node values were taken as robust if they were recov-
ered with support values of UFBS $95 and SH-aLRT $80
(Guindon et al., 2010;Bui et al., 2013). Substitution models
were selected using the Bayesian information criterion im-
plemented in ModelFinder (Kalyaanamoorthy et al., 2017).
Uncorrected p-distance was calculated for the new species
and comparative species of Paracheilinus using GENEIOUS
Prime. We used median-joining (Bandelt et al., 1999) in
PopART (Leigh and Bryant, 2015) to construct haplotype
networks for select species of Paracheilinus based on a subset
of the 642 mitochondrial COI alignment described above.
Taxonomy
Infraclass Teleostei
Order Labriformes
Family Labridae
Genus Paracheilinus Fourmanoir, 1955
Paracheilinus Fourmanoir in Roux-Este
`ve and Fourmanoir,
1955:100(typespeciesParacheilinus octotaenia Fourmanoir,
by original designation).
Diagnosis.—The following synapomorphy and combination
of characters diagnose Paracheilinus from all other labrid
genera.
Shape of pelvic girdle:Paracheilinus possesses a distinct
pelvic girdle characterized by anterior narrowing of the cen-
tral processes with the external ventral wings curling medi-
ally along the mid-line. This curling forms a shallow canal
that extends along the anterior half to two-thirds the length
of the pelvic girdle (described and illustrated in Tea et al.,
2021). We consider this character a synapomorphy diagnos-
ing Paracheilinus.
In addition to the aforementioned character, the follow-
ing combination of additional characters distinguishes Par-
acheilinus from all other labrid genera: dorsal-fin rays IX
(rarely VIII or X),11; anal-fin rays III,9 (rarely 8 or 10); pec-
toral-fin rays 14 (rarely 13 or 15); principal caudal-fin rays
13, the median 11 branched; lateral line interrupted, pored
scales in the dorso-anterior series 11–17, pored scales in the
midlateral posterior peduncular series 3–10; median predor-
sal scales 5 (rarely 4 or 6); rows of scales on cheek 2; total
gill rakers 11–18; branchiostegal rays 5; vertebrate 9 þ16;
scleral cornea of orbit over pupil bisected into two foci; lips
small, without plicate folds, barely visible when mouth
closed; three pairs of recurved canines anteriorly on upper
jaw, third pair largest; single pair of canines anteriorly on
lower jaw; sides of jaw with single row of small, closely set
conical teeth; no canines at corner of mouth; palate with-
out teeth; ventral margin and corner of preopercle thin and
membranous, posterior bony edge smooth (sometimes very
weakly serrated); snout, chin, and interorbital space naked;
pelvic fin short, longest ray not reaching anal-fin origin;
dorsal scleral surface of orbit and orbital rim of juveniles
and initial phase (IP) specimens with white “eyebrow”
markings.
Description.—Dorsal-fin rays IX (rarely VIII or X),11, seg-
mented rays variably branched (see below); anal-fin rays
III,9 (rarely 8 or 10), segmented rays variably branched (see
below); pectoral-fin rays 14 (rarely 13 or 15), upper two un-
branched; pelvic-fin rays I,5; principal caudal-fin rays 7 þ6,
uppermost and lowermost unbranched; upper procurrent
caudal-fin rays 4–5; lower procurrent caudal-fin rays 4–6;
lateral line interrupted, with dorso-anterior series of pored
scales 11–17 and midlateral posterior peduncular series 3–
10; first pored scale on posterior peduncular series often pit-
ted; last pored scale on posterior peduncular series enlarged
and overlapping hypural crease; scales above lateral line to
origin of dorsal fin 2; scales below lateral line to origin of
anal fin 6; median predorsal scales 5 (rarely 4 or 6); circum-
peduncular scales 14–16; total gill rakers 11–18; branchios-
tegal rays 5; vertebrae 9 þ16.
Mouth small, oblique, maxilla not reaching vertical at an-
terior edge of orbit; dentition typical of genus (see diagnosis);
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lips small, without plicate folds, barely visible when mouth
closed; gill rakers short, longest about a third length of lon-
gest filament on first gill arch; posterior nostril an oval open-
ing about 2–3 times larger than cephalic sensory pores, about
level with fleshy upper edge of orbit and slightly anterior to
vertical at anterior bony rim of orbit; anterior nostril smaller,
with a short fleshy rim anterior and slightly ventral to poste-
rior nostril.
Head scaled except for anterior occipital region, interorbit-
al space, snout, and chin; a row of elongate scales on dorsal-
and anal-fin bases; basal half of caudal fin with large scales;
axillary scale of pelvic fin slightly shorter than pelvic spine;
midventral scaly process of pelvic fins slightly longer than
pelvic spine; free ventral margin of preopercle extending for-
ward to vertical at center of orbit, vertical posterior margin to
level of lower edge of pupil; exposed bony edge of preopercle
smooth without serrations (sometimes very weakly serrated
on ventral bony edge).
Origin of dorsal fin above third lateral-line scale; dorsal-
fin spines progressively longer; posterior soft-portion of dor-
sal fin with or without filaments, branching pattern variable
(see below); origin of anal fin below base of first dorsal-fin
soft ray; pelvic fins short, longest ray barely reaching anal-
fin origin; caudal fin varying from truncate, rounded, emar-
ginate (with or without lobes bearing filamentous exten-
sions), or lanceolate (in one species).
Variable branching of dorsal- and anal-fin rays.—Paracheilinus
is one of several labrid genera to possess filamentous dorsal-
fin rays. The term “filament” is loosely defined, but generally
refers to any filamentous prolongation of spinous or radial
elements in the fins. In most filament-bearing labrid gen-
era, those borne on the dorsal fin occur only in the unseg-
mented, spinous portion of the fin. These are, however,
usually fibrous extensions of the cirri or flexible extensions
of membranous tissue distal to the spinous element. Para-
cheilinus is unusual in having dorsal-fin filaments only in the
segmented rays in the posterior portion of the fin. In Paracheili-
nus,thefilamentsarealwaysprolongationsoftheradialele-
ments and not fibrous or membranous extensions. Filaments
can consist of a single segmented ray, or multiple adjacent rays
bounded by membranous tissue. In the former condition, it is
not uncommon for several species to develop more than one
filament, but the filaments are usually separated from each
other by deep incisions of the interradial membrane. The fila-
ments in some species can be very long (up to 1.8 in SL in
P. nursalim) and make for exceptionally spectacular displays
when fully raised.
Within the genus, the following western Indian Ocean spe-
cies form a monophyletic sister group to all remaining spe-
cies of Paracheilinus. They are P. attenuatus,P. octotaenia,P.
hemitaeniatus, and P. piscilineatus. This relationship is sup-
ported based on molecular sequence data (Allen et al., 2016).
Except for P. attenuatus, which is unusual in possessing a sin-
gle hair-like “naked” ray with no surrounding membranous
tissue, the remaining species lack filaments entirely. No other
cirrhilabrin genera possess segmented dorsal-fin filaments.
The character therefore appears to be a derived condition
unique to Paracheilinus, but not a synapomorphy diagnosing
it. Given that there are several other non-filament bearing
species nested within the genus, we consider the presence of
segmented dorsal-fin filaments homoplasious within Para-
cheilinus. The only other labrids to somewhat approach this
morphological condition are those in the hypsigenyin genera
Bodianus,Clepticus,andSemicossyphus,wherelargemales
may develop pronounced or slightly filamentous seg-
mented dorsal-fin rays, though never as dramatically as in
Paracheilinus.
The degree of segmented dorsal-fin ray branching is fur-
ther unusual in Paracheilinus and is highly variable within
and between species. For species that possess filaments, the
filamentous rays are usually unbranched, but those situated
more posteriorly (in species bearing more than one fila-
ment) may exhibit weak primary or, rarely, secondary and
tertiary branching. The last segmented dorsal-fin ray is al-
ways split to base regardless. If branching occurs in the fila-
mentous rays, it is usually near the base, with the branched
portions rarely extending the full length of the filament.
This condition does not appear to be ontogenetic, as juve-
niles and IP males can also display variable branching of
segmented dorsal-fin rays.
For species that lack filaments, segmented rays exhibit
branching patterns typical of other labrids. In Paracheili-
nus, however, it is not uncommon for some of the anterior
segmented rays to be completely unbranched, the number
and positioning of which is variable. For example, in the
holotype of the non-filament bearing P. t o g e a n s i s , only the
first segmented ray is unbranched, followed by branching
of all proceeding rays (Kuiter and Allen, 1999). Underwater
images of three separate individuals of P. togeansis published
in Allen et al. (2016) demonstrate unbranching of the ante-
riormost three, four, and five segmented rays, respectively
(Allen et al., 2016: figs. 49–50). An unpublished photograph
(examined by the authors of this study) of P. togeansis pho-
tographed by Ned DeLoach shows yet a specimen with
branching of all segmented dorsal-fin rays. More uncom-
monly, unbranched rays may occur in between branched
rays (Allen et al., 2016: fig. 52). Rarely, filaments may de-
velop on the anal fin, though the presence and number of
anal-fin filaments are aberrant and are never diagnostic for
Fig. 2. Paracheilinus paineorum (non-Australian species) demonstrat-
ing filamentous prolongation of segmented dorsal- and anal-fin rays.
In this specimen, nearly all dorsal-fin rays are unbranched. Note that
the bifurcation at the tip of the first dorsal filament is not supported by
branching of the segmented ray (it appears to be a result of a tear in
the surrounding membrane). Note the presence of aberrant anal-fin fil-
aments. Underwater photograph taken in Lembeh Strait, Indonesia, by
N. DeLoach.
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any of the known species (Fig. 2). These conditions become
increasingly prevalent in hybrids (see fig. 5B in Allen et al.,
2016).
While none of these conditions in isolation are unique
among teleost fishes, the degree of variation displayed in
ray branching pattern within a single genus is unusual. No
other labrid genus displays such variability in the position-
ing, number, and extent of branched dorsal- and anal-fin
rays in any given species. Given the labile nature of these
characters, however, we do not at this time consider this a
synapomorphy diagnosing the genus, though we acknowl-
edge its potential usefulness in separating certain species
from other labrid genera.
Etymology.—Combination of the Greek para meaning near
or proximal to, and the labrid genus Cheilinus, in reference
to its superficial resemblance to the genera Cheilinus and
Pseudocheilinus (Roux-Este
`ve, 1956); to the former in having
the second anal-fin spine shorter than the third, and to the
latter in having recurved outward pointing canines. Para-
cheilinus is, however, most closely related to Cirrhilabrus.
Genus is masculine. Species of Paracheilinus are commonly
known as flasher wrasses after their spectacular courtship
displays.
Habitat and biology.—Small to medium (usually under 70 mm
SL) brightly colored labrids frequently found swimming above
rubble pans with Halimeda or Padina growth, in groups consist-
ing anywhere between tens to hundreds of individuals with
females greatly outnumbering males. Occurs between 5–70 m,
but with most species inhabiting 15–40 m (Allen et al., 2016).
Diet primarily zooplankton which they pick off the water col-
umn. Males of all species are known for flashing bright irides-
cent colors during acts of aggression or nuptial display. This
display is often accompanied by rapid swimming and the erec-
tion of all median fins. When not displaying, the colors are
muted (though still very striking) and the fins are folded against
the body.
KEY TO THE SPECIES OF AUSTRALIAN PARACHEILINUS
1a. Body with stripe pattern A or A’; caudal fin truncate to
slightly emarginate and with pronounced filamentous
lobes; dorsal fin with four or more yellow to red-
dish orange filaments ------------------------------- ---------------------------------------------------------------------------- -- 2
1b. Body with stripe pattern B; caudal fin rounded and
without filaments; dorsal fin with three or fewer
red filaments (usually one) ------------------------------------------------------------------------------------------------------- 3
2a. Body with stripe pattern A; ground color of body
red to maroon in life; dorsal fin and filaments red-
dish orange at rest, richly yellow when in display;
central portion of caudal fin with triangular hya-
line region, upper and lower lobes red with metallic
blue margins dorsally and ventrally (in life); spines
and rays of median fins purple in preservation
(northern Great Barrier Reef)---------------------------- P. filamentosus
2b. Body with stripe pattern A’; body with two black
rectangular epaulettes, one below spinous portion
of dorsal fin, one along lower caudal peduncle
(dusky in preservation); ground color of body yel-
low orange in life; dorsal fin and filaments yellow
at rest, filaments bright white when in display; cen-
tral portion of caudal fin with intricate blue mark-
ings (in life); spines and rays of median fins
translucent and without purple pigment in preser-
vation (Timor Sea) --------------------------------------------------------------------------------------- P. nursalim
3a. Anal fin uniformly yellow or orangey yellow; dorsal
fin with one to four filaments (usually one; Timor
Sea)--------------------------------------------------------------------------------------------------------------------------------------------- P. flavianalis
3b. Anal fin sharply bicolored, yellow basally, red dis-
tally; dorsal fin with one filament (Coral Sea and
Great Barrier Reef) ------------------------------- P. amanda, new species
Paracheilinus amanda, new species
urn:lsid:zoobank.org:act:84FE0313-6FEF-43EC-9DB1-
DE52090AB295
Amanda’s Flasher Wrasse
Figures 3,4,5A1–A2,6,7;Tables 1–2
Paracheilinus rubricaudalis (non Randall and Allen, 2003):
Allen et al., 2016: 73, fig. 47 (aquarium specimen from
Great Barrier Reef; image reproduced here [Fig. 2]); also
misidentified as P. rubricaudalis in molecular phylogeny.
Holotype.—QM I.39758, 47.6 mm SL, Harrier Reef, Great
Barrier Reef, 158080S, 1458410E, 20 m, T. Bennett, 27 October
2013.
Fig. 3. Paracheilinus amanda, new species, aquarium specimen from
Harrier Reef, the Great Barrier Reef. Specimen not retained. Photograph
by K. Endoh. Fig. 4. X-ray of Paracheilinus amanda,newspecies,AMSI.50116-001,
paratype, showing aberrant branching of lowermost principal caudal ray.
402 Ichthyology & Herpetology 111, No. 3, 2023
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Paratypes.—AMS I.50116-001, 44.4 mm SL, Flora Reef, Coral
Sea, 168460S, 1478460E, 42 m, hand nets, T. Bennett, 2 Febru-
ary 2022 (GenBank accession number OQ605966); WAM
P.33973.001, 2, 37.3–43.7 mm SL, Harrier Reef, Great Barrier
Reef, 158080S, 1458410E, 35–36 m, hand nets, T. Bennett, 1
September 2013; ZRC 64175, 2, 32.3–47.6 mm SL, 40–50 m,
hand nets, G. Gaylan, J. M. Nugas, and J. Genoso, off Hula,
southern Papua New Guinea, Coral Sea, 28 March 2023.
Diagnosis.—Dorsal-fin rays IX,11; anal-fin rays III,9; pecto-
ral-fin rays 14; pelvic-fin rays I,5; pored lateral-line scales 14–
17 þ5–7 ¼19–24; gill rakers 5–6 þ11–12 ¼16–18; body
Fig. 5. A selection of Paracheilinus in life. (A1) P. amanda, new species, ZRC 64175, male paratype, 47.6 mm SL, off Hula, southern Papua New
Guinea, Coral Sea; (A2) P. amanda, new species, underwater photograph from Osprey Reef, Coral Sea; (B) P. carpenteri, underwater photograph
from Mabini, Batangas, Philippines. Note the darkened posterior dorsal- and caudal-fin bases and the presence of a second stripe behind the pec-
toral fin; (C) P. flavianalis, underwater photograph from Bali, Indonesia; (D) P. mccoskeri, underwater photograph from Khao Lak, Thailand; (E) P.
rubricaudalis, underwater photograph from Mborokua, Solomon Islands. Note the reduced markings on caudal fin. Photographs by H. H. Tan (A1);
T. Yamazumi (A2); T. Cameron (B); V. Chalias (C); T. Kawamoto (D); and N. DeLoach (E).
Tea and Walsh—Review of Australian Paracheilinus with description of a new species 403
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depth 3.2–3.8 in SL; head length (HL) 3.1–3.3 in SL; snout
length 4.0 in HL; orbit diameter 3.0–3.6 in HL; interorbital
width 3.0–3.4 in HL; least depth of caudal peduncle 2.3 in HL;
caudal peduncle length 1.7–2.3 in HL; TP males with one elon-
gate, red filamentous dorsal-fin ray, longest dorsal-fin soft ray
2.1–5.5 in SL; pelvic fin length 2.0–2.8 in HL; caudal fin of TP
males round without filamentous lobes; body with stripe pat-
tern B; basal third of anal fin bright yellow, remaining distal
two-thirds orange red, demarcation of both colors with in-
complete row of blue spots (usually completely absent).
Description.—Dorsal-fin rays IX,11; anal-fin rays III,9; last
dorsal- and anal-fin rays split to base; pectoral-fin rays 14,
upper two unbranched; pelvic-fin rays I,5; principal caudal-
fin rays 7 þ6, uppermost and lowermost unbranched (low-
ermost principal caudal-fin ray aberrantly branched in AMS
I.50116-001; Fig. 4); upper procurrent caudal-fin rays 5;
lower procurrent caudal-fin rays 5 (4–5); lateral line inter-
rupted, with dorsoanterior series of pored scales 17 (14–17)
and midlateral posterior peduncular series 6 (5–7); first
pored scale on posterior peduncular series often pitted; last
pored scale on posterior peduncular series enlarged and
overlapping hypural crease; scales above lateral line to ori-
gin of dorsal fin 2; scales below lateral line to origin of anal
fin 6; median predorsal scales 5; median preventral scales 5
(3–6); transverse scale rows on cheek 2; circumpeduncular
scales 16 (14–16); gill rakers 6 (5–6) þ11 (11–12) ¼17 (16–
18); vertebrae 9 þ16 (Fig. 4).
Body depth 3.2 (3.2–3.8) in SL; body width 2.1 (1.9–2.3) in
body depth; head length 3.1 (2.8–3.3) in SL; snout length 4.0
(4.0–4.4) in HL; orbit diameter 3.6 (3.0–4.5) in HL; interorbital
width 3.4 (3.0–4.5) in HL; least depth of caudal peduncle 2.3
(2.3–2.9) in HL; caudal-peduncle length 1.8 (1.7–2.3) in HL.
Mouth small, oblique, maxilla not reaching vertical at
front edge of orbit, upper jaw 4.0 (3.9–6.4) in HL; three pairs
of curved canine teeth anteriorly in upper jaw, progressively
more laterally projecting, third (posteriormost) pair largest;
single pair of canine teeth anteriorly in lower jaw, very
strongly curved laterally; side of jaws with single row of small
close-set conical teeth; no canine tooth at corner of mouth;
no teeth on palate; fleshy flap on side of lower lip; gill rakers
short, longest about one-third length of longest gill filaments
on first gill arch; posterior nostril an oval opening about 2–3
times larger than cephalic sensory pores, about level with
A
B
C
C
C
C
P. a m a n d a
P. c a r p e nt e r i
P. f l a v i an a l i s
P. m c c o s ke r i
P. r u b r i ca u d a l i s
P. f i l a m en t o s u s
P. n u r s a li m
P. mccoskeri KF709098
P. rubricaudalis OQ600404
P. flavianalis KF709105
P. rubricaudalis OQ600405
P. carpenteri KP187641
P. flavianalis OQ600543
P. rubricaudalis OQ600403
P. flavianalis KP187637
P. carpenteri KP187642
P. mccoskeri KP187624
P. carpenteri KP187639
P. rubricaudalis OQ600406
P. amanda sp. nov. KT253628
P. flavianalis KP187631
P. mccoskeri KF709099
P. flavianalis KF709103
P. mccoskeri KP187625
P. carpenteri KF709101
P. flavianalis KP187630
69.7/89
9
7.4/
99
9
2.4/
93
8
7.9/97
77.3/89
9
7.2/
99
0
.01
s
u
bs
/
s
it
e
KT253628
P. amanda sp. nov. OQ605966
KP187639
KF709101
KP187642
KP187641
KP187624
KP187625
KF709099
KF709098
KP187630
KF709105
KF709103
KP1876387
OQ600543
KP187631
OQ600403
OQ600404
OQ600405
OQ600406
KT253628
OQ605966
Fig. 6. (A) Maximum-likelihood phylogenetic relationships for select species of Paracheilinus inferred using mitochondrial COI. Support values at
nodes correspond to SH-aLRT and UFBS support values, respectively. Scale bar indicates the number of substitutions per site. Outgroup taxa were
pruned from the tree (see Allen et al., 2016). See Data Accessibility for tree file. (B) Median-joining haplotype network inferred using mitochondrial
COI. Each circle represents a haplotype, and its size is proportional to its total frequency. Each black crossbar represents a single nucleotide change.
(C) Geographical distribution for select species of Paracheilinus. Members of the P. mccoskeri species complex are represented by colored circles.
Distribution records for P. carpenteri and P. mccoskeri are not shown in full—the former extends into southern Japan (indicated by the green bro-
ken arrow), and the latter extends into the western Indian Ocean (indicated by the purple broken arrow). Bicolored symbols indicate sympatry in
that locality. Colored stars indicate distribution records for P. filamentosus and P. nursalim in Australia.
404 Ichthyology & Herpetology 111, No. 3, 2023
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fleshy upper edge of orbit and slightly anterior to vertical at
anterior bony rim of orbit; anterior nostril smaller, with a
short fleshy rim anterior and slightly ventral to posterior nos-
tril; internarial space about 1.0 in orbit diameter.
Head scaled except for interorbital space, snout, and chin;
a row of pointed scales on base of dorsal and anal fins; basal
half of caudal fin with large scales; axillary scale of pelvic fin
slightly longer than pelvic spine; midventral scaly process
Fig. 7. Paracheilinus amanda, new species (above right), aquarium specimen from Harrier Reef, Great Barrier Reef; P. rubricaudalis (middle right)
underwater photograph from Solomon Islands; and P. flavianalis (below right), underwater photograph from Bali. The purpling of median-fin rays
in preserved specimens are shown in the lettered insets. (A) Paracheilinus amanda, QM I.39758, holotype; (B–C) Paracheilinus rubricaudalis,
WAM P.33979.001; (D) Paracheilinus flavianalis, NTM S.18121-011. Live photographs of P. amanda,P. rubricaudalis, and P. flavianalis by F.
Walsh, D. Brown, and K. Nishiyama, respectively. Live specimens not retained. Photographs of preserved specimens by T. Miskiewicz and Y. K. Tea.
Tea and Walsh—Review of Australian Paracheilinus with description of a new species 405
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Table 2. Summary of live coloration characters for select species of Paracheilinus. Unless specified, all characters apply only to TP males.
Paracheilinus amanda,
new species Paracheilinus carpenteri Paracheilinus flavianalis Paracheilinus mccoskeri Paracheilinus rubricaudalis
Body coloration Pattern B Pattern B, but with an addi-
tional stripe behind pecto-
ral fin
Pattern B Pattern B Pattern B
Dorsal-fin filament Always one; always red 2–6 (usually 3–4); yellow to
red
1–4 (usually one); always red Always one; always yellow Always one; yellow to red
Posterior dorsal fin Ground color yellow hyaline
with extensive metallic
blue markings
Ground color yellow to red
base darkened with elon-
gate black blotch
Ground color yellow hyaline
with metallic blue markings
on distalmost edge
Ground color yellow with ex-
tensive metallic blue
markings
Bright red without any blue
markings
Caudal fin Typical of pattern B Typical of pattern B, but base
darkened with black
Typical of pattern B Typical of pattern B Typical of pattern B, but
bright red and with the
concentric bands
attenuated or absent
completely
Anal fin Bicolored; basal third bright
yellow, distal two-thirds
bright orange red. Blue
spots absent or incomplete
when present.
Bicolored; basal third bright
yellow, distal two-thirds
bright orange red. Blue
spots usually present,
sometimes absent entirely.
Uniform yellow to yellowish
orange, rarely suffused with
red on the outer edge.
Blue spots usually present,
sometimes absent entirely.
Bicolored; basal third bright
yellow, distal two-thirds
bright orange red. Blue
spots usually present,
sometimes absent entirely.
Interradial membrane rich
orange yellow. Anal-fin rays
bright purple red.
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of pelvic fins slightly shorter than pelvic spine. Free ventral
margin of preopercle extending to vertical at center of orbit,
vertical posterior margin to level of lower edge of orbit; ex-
posed bony edge of preopercle smooth without serrations.
Origin of dorsal fin above third lateral-line scale, predorsal
length 3.2 (2.8–3.2) in SL; dorsal-fin spines progressively
longer, first 9.6 (9.2–14.6) in HL, and ninth 2.1 (2.0–2.7) in
HL; one to three elongate filamentous segmented rays on
posterior dorsal fin in males, bound together by membrane,
first longest, 2.1 (1.8–5.7) in SL; origin of anal fin below
base of last dorsal-fin spine, preanal length 1.9 (1.7–1.9) in
SL; first anal-fin spine 5.1 (4.5–9.6) in HL; second anal-fin
spine 3.8 (3.6–5.0) in HL; third anal-fin spine 3.0 (3.0–4.3)
in HL; longest anal-fin soft ray 4.2 (3.7–8.1) in SL; caudal fin
rounded, 3.6 (3.6–4.6) in SL; pectoral-fin length 1.6 (1.5–
1.7) in HL; pelvic-fin length 2.0 (2.0–2.8) in HL.
Coloration of males in life.—Based on color photographs of
specimens when freshly dead, and live individuals photo-
graphed in aquaria (Figs. 3,5A1–A2,7): head and body
orange to orange brown with body stripes following pattern
B; stripes purplish blue to bright neon blue in life; iris bright
orange with bright yellow ring around pupil; spinous por-
tion of dorsal fin yellow; posterior soft dorsal fin hyaline
overlain with bright metallic blue, sometimes breaking into
indistinct spot-bands medially and on distal edge; first two
to three interradial membrane spaces (including the fila-
ment) bright red; basal third of anal fin bright yellow, re-
mainder distal portion of fin bright orange red, interphase of
both colors usually without blue spots, or if present, usually
incomplete and towards posterior portion of fin; distalmost
edge of anal fin bright blue; caudal fin hyaline with a pair of
concentric blue bands following contour of fin, first on basal
third, second on outermost edge; pelvic fins reddish hyaline,
bright blue on distalmost edge; pectoral fins hyaline.
Coloration of females in life.—Females typical of species in
this species complex. Head and body orange to orange
brown with body stripes following pattern B; stripes pur-
plish blue to neon blue in life. Median fins yellow hyaline
with indistinct blue markings.
Color in preservation.—Uniformly pale tan, except body
stripe pattern now dark tan. All fins translucent hyaline.
Segmented fin rays on median fins in the largest specimen
(47.6 mm SL holotype) purple.
Habitat and distribution.—Paracheilinus amanda is known
from Harrier Reef in the northern Great Barrier Reefs, as well
as Flora, Holmes, and Osprey Reefs in the Coral Sea. It is
also known from specimens collected off Taurama and Hula
in southern Papua New Guinea, along the north-western
margin of the Coral Sea. The identity and collection locality
of the specimen of P. rubricaudalis used in the molecular
phylogeny of Allen et al. (2016) is erroneous. The specimen
is P. amanda, collected from Harrier Reef, not Ribbon Reef,
alongside the WAM paratypes (WAM P.33973.001). The spe-
cies frequents habitat typical of the genus, consisting of
low-lying rubble pans between 20 to 50 m. It is replaced by
the closely related P. carpenteri in Japan, Philippines, Brunei,
eastern Sulawesi, and Palau, P. flavianalis in Indonesia (Bali
eastwards in the Lesser Sunda Islands and the Moluccas),
Timor Leste, West Papua, and north-western Australia,
P. mccoskeri in the Indian Ocean, and P. rubricaudalis in
wider Melanesia (northern Papua New Guinea, the Solomon
Islands, Fiji, and Vanuatu).
Etymology.—The species is named in honor of Amanda Hay,
ichthyology collections manager at the Australian Museum,
Sydney. With over 25 years of experience in ichthyological
collections and research, she has not only contributed sig-
nificantly towards the study of Australasian fishes, but
also supported and assisted the research endeavors of
many ichthyologists of all career stages working at the
Australian Museum. The name amanda is treated as a
noun in apposition.
Comparisons and phylogenetic interpretation.—Paracheilinus
amanda most closely resembles P. carpenteri,P. flavianalis,P.
mccoskeri, and P. rubricaudalis (Fig. 5). The five species form a
complex of largely allopatric species united in sharing stripe
pattern B, hereafter referred to as the P.mccoskeri complex.
Except for P. carpenteri and P. flavianalis, members of the P.
mccoskeri complex typically possess a single dorsal-fin fila-
ment through prolongation of the anteriormost 1–3 seg-
mented rays. Since all five species possess overlapping
morphometric and meristic characters, separation of species
is most reliable through comparison of live coloration of TP
males, which typically involves a combination of coloration
details of the dorsal filament(s), posterior dorsal fin, and
anal fin. As with many species of Paracheilinus, intraspecific
variation is typically very high, although some trends are
evident within each species. This variation is further exacer-
bated by the proclivity for hybridization displayed in several
species. As such, it is possible to encounter individuals that
depart from the following character summaries.
In P. amanda (Figs. 3,5A1–A2,7), the anal fin is sharply bi-
colored, with the basal third yellow and the distal region
bright orange red. The interphase of both colors on the anal
fin is incompletely lined with bright blue spots (spots usu-
ally absent entirely). The dorsal fin is always with a single
red filament, and the posterior portion of the dorsal fin is
extensively decorated in metallic blue. The segmented rays
of the median fins are deep purplish red in life, turning
purple in alcohol.
In P. carpenteri (Fig. 5B), the anal fin is usually sharply bi-
colored (as with P. amanda) and usually, but not always,
with a complete series of blue spots along the colored inter-
phase. The dorsal fin is ornamented with 2–6 (usually 3–4)
filaments that vary in color from yellow to red. The bases of
the caudal fin and posterior dorsal fin are often blackened
(the latter with an elongate black blotch). In addition to dis-
playing the typical assortment of stripes in pattern B, there
is always an additional short stripe behind the pectoral fin.
Large males can sometimes have the body stripes broken
and somewhat reticulate.
In the highly variable P. flavianalis (Figs. 5C,7,8,9), the
anal fin is never sharply bicolored, and is almost always uni-
formly yellow or orangey yellow (very rarely it may be
washed with red near the distal edge, but this demarcation
is always suffused; Fig. 8B). There is usually, but not always,
a complete series of blue spots (cf individuals in Figs. 5C,7,
8B, D) on the anal fin. The dorsal fin is ornamented with 1–
4 (usually one) red filament(s). The segmented rays of the
median fins turn purple in alcohol.
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In P. m c c o s k e r i (Fig. 5D), the anal fin is usually, but not
always, sharply bicolored (as with P. a m a n d a and P. c a r p e n -
teri), with the basal third yellow and the distal region
bright orange red. Occasionally, individuals may have en-
tirely yellow or red anal fins (the latter phenotype is more
common in the western Indian Ocean). The interphase of
both colors on the anal fin is usually, but not always,
completely lined with bright blue spots. The dorsal fin is
always with a single yellow filament, and the posterior
dorsal fin is richly decorated in metallic blue on the distal
edge.
In P. rubricaudalis (Figs. 5E,7), the interradial membranes
of the anal fin are a rich orange yellow that contrast strong-
ly with the bright purplish red segmented rays, giving the
appearance of deep serrations on the fin. The anal fin is al-
most always without blue spots. The dorsal fin is always
with a single yellow or red filament, and the posterior dorsal
fin is bright red without metallic blue markings. The caudal
fin is bright red and with the usual concentric bands attenu-
ated or completely absent.
Among members of the P. m c c o s k e r i complex, P. rubricauda-
lis,P. amanda,andP. flavianalis share the unusual property of
having the segmented rays on their median fins turning pur-
ple in alcohol (Fig. 7). In contrast, the rays of P. m c c o s k e r i and
P. carpenteri are either translucent, or in large individuals,
weakly blue green, in preservation. The unusual quality of
purple fin rays and other osseous elements was briefly re-
viewed by Tea et al. (2022a) for species of Cirrhilabrus,butthe
condition has not been extensively reviewed for Paracheilinus
and other labrid taxa. Several other species of Paracheilinus de-
velop purpling of radial elements in alcohol (see Allen et al.,
2016), including P. filamentosus (see below). Character sum-
maries for all species of the P. m c c o s k e r i complex are presented
in Table 2.
Fig. 8. Select individuals of Paracheilinus flavianalis demonstrating variability in the number of dorsal-fin filaments, coloration of anal fin, and
spot band pattern on the anal fin. (A) Underwater photograph from Triton Bay, Indonesia; (B) underwater photograph from Wakatobi, Sulawesi,
Indonesia; (C–D) underwater photographs from Bali, Indonesia. Photographs by R. Smith (A); J. Castellano (B); W. Osborn (C); and R. H. Kuiter (D).
Fig. 9. Paracheilinus flavianalis, underwater photograph from Hibernia
Reef, Timor Sea, north-western shelf of Western Australia. Note sympat-
ric Cirrhilabrus morrisoni and habitat consisting of extensive Halimeda
growth. Photograph by G. R. Allen.
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Molecular phylogenetic analyses of mitochondrial COI
recover the P.mccoskeri complex as monophyletic, suggest-
ing that pattern B is synapomorphic for this complex and
apomorphic within Paracheilinus (Fig. 6A). Paracheilinus
amanda,P. f l a v i a n a l i s ,andP. rubricaudalis form a monophy-
letic lineage within the P. mccoskeri complex, with the
three sharing successive sister relationships to P. c a r p e n t e r i
and P. mccoskeri (Fig. 6A). This relationship is supported by
a single morphological character (i.e., purpling of radial el-
ements) and is mostly congruent to those described in
Allen et al. (2016), except that in their study, P. rubricauda-
lis was not included in the analyses; the P. r u b r i c a u d a l i s in
Allen et al. (2016) is P. a m a n d a (GenBank accession number
KT253628).
While the haplotype diversity for each species corre-
sponds well with species-specific coloration pattern and
geographical distribution (Fig. 6B), phylogenetic resolu-
tion based on mitochondrial COI was unsatisfactory for
the lineage comprising P. a m a n d a ,P. ru b r i c a u d a l i s , and
P. flavianalis.Paracheilinus amanda and P. r u br i c a u d a l i s were
recovered as monophyletic sister lineages, but the relation-
ship of P. flavianalis could not be resolved due to low se-
quence variability in mitochondrial COI. This scenario is
not uncommon for groups of coral reef fishes where sexual
selection favors female mate choice and male coloration,
leading to incongruent signals in morphological and mo-
lecular data, or those that have speciated only very recent-
ly (Victor and Randall, 2014;Hench et al., 2019). Pairwise
comparison of mitochondrial COI reveals a genetic dis-
tance of 1–1.2% (uncorrected p-distance) between P.
amanda and P. r u b r i c a u d a l i s ,andbetweenP. a m a n d a and P.
flavianalis.GeneticdistancesbetweenP. r u b r i c a u d a l i s and
P. f l a v i an a l i s were less than 1%, suggesting that separation
of both species occurred only very recently. All three species
are allopatric (Fig. 6C). Notwithstanding the lack of phyloge-
netic resolution, P. a m a n d a is at the very least distinct from all
congeneric species on the basis of morphology and molecular
sequence data.
Remarks.—The paratypes of the new species consist most-
ly of non-TP males, which may account for the larger var-
iation in dorsal-fin filament lengths when compared
with other members of its species complex (17.6–54.4%
SL vs. 27.8–50.3% SL in our examined specimens of P. f l a -
vianalis;and45.8–64.1%SLinourexaminedspecimens
of P. r u b r i c a u d a l i s ). Live photographs of TP males and
morphometric data from the two TP males in the type se-
ries (Figs. 3,5A1,5A2,7), however, show proportionately
longer dorsal-fin filaments in P. a m a n d a compared to
lengths typically displayed by related species. The width
of the dorsal-fin filament appears to be broadest in
P. a m a n d a ,consistingofthreesegmentedrays(vs.oneto
two) at the base of the filament in all specimens exam-
ined in the type series. More comparative material is
needed to determine if this character is restricted to this
species.
The AMS paratype of P. a m a n d a (AMS I.50116-001) has
the lowermost principal caudal-fin ray deeply bifurcate
and branched close to the base (Fig. 4). An X-ray of the
specimen suggests that this branching is aberrant and like-
ly a result of improper healing of an injury. The ray imme-
diately above appears to have been injured in the same
position.
Paracheilinus filamentosus Allen, 1974
Filamentous Flasher Wrasse
Figures 10,11A–E
Paracheilinus filamentosus Allen, 1974: 452 (holotype AMS
I.16994-001, Kranket Island, Madang, Papua New
Guinea); Randall and Lubbock, 1981: 26, pl. 2C (gener-
ic revision, specimens from Papua New Guinea and the
Solomon Islands [BPBM 16007], but not those from
Indonesia and the Philippines); Kuiter and Tonozuka,
2004: 494 (field guide, Papua New Guinea and the
Solomon Islands, but not those from Indonesia and
Palau); Michael, 2009: 284 (field guide, underwater
photograph from Papua New Guinea, but not those
from Indonesia, Palau, the Philippines); Kuiter, 2010:
106, figs. B, C (underwater photographs from Papua
New Guinea and the Solomon Islands, but not those
from Indonesia); Allen and Erdmann, 2012:706(field
guide, Papua New Guinea and the Solomon Islands,
but not those from Brunei, Indonesia, Palau, the
Philippines, and West Papua).
Diagnosis.—Dorsal-fin rays IX,11 (holotype VIII,11); anal-
fin rays III,9; pectoral-fin rays 14; pelvic-fin rays I,5; pored
lateral-line scales 14–17 þ5–10 ¼19–27; gill rakers 5–6 þ
7–10 ¼12–16; body depth 3.0–3.5 in SL; head length 2.9–
3.5 in SL; snout length 3.7–4.8 in HL; orbit diameter 3.1–
4.9 in HL; interorbital width 3.4–4.6 in HL; least depth of
caudal peduncle 1.9–2.5 in HL; caudal peduncle length
1.6–2.8 in HL; TP males with 4–7 elongate filamentous dor-
sal-fin rays, longest dorsal-fin soft ray 1.7–2.0 in SL; pelvic
fin length 1.9–2.1 in HL; caudal fin of TP males with pro-
nounced filamentous lobes; caudal-fin length 3.7–5.0 in
SL; caudal concavity 2.3–6.4 in SL; body with stripe pat-
tern A; dorsal fin reddish orange at rest, richly yellow
when in display; ground color of body red to maroon in
life; central portion of caudal fin with triangular hyaline
region, upper and lower lobes red with metallic blue mar-
gins dorsally and ventrally (in life); spines and rays of me-
dian fins purple in preservation.
Remarks.—Allen (1979) described P. filamentosus on the basis
of 23 specimens collected from Papua New Guinea and the
Solomon Islands. He noted the holotype as having VIII,11
dorsal-fin rays (versus the usual IX,11) for the genus. An
Fig. 10. Paracheilinus filamentosus, AMS I.40666-002, 46.6 mm SL,
freshly euthanized male, Lizard Island, northern Great Barrier Reef,
Australia. Photograph by J. Leis.
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additional seven specimens from Saparua, Moluccas, Indo-
nesia, were designated as paratypes (USNM 209923) in an
addendum published at the end of the original description.
Subsequent work published by other authors expanded the
known distribution of P. fi l a m e n t o s u s to include Brunei,
various localities across Indonesia, Palau, and the Philip-
pines. In the most recent revision of the genus, Allen et al.
(2016) restricted P. f i l a m e n t o s u s to Papua New Guinea and
Fig. 11. Paracheilinus filamentosus, images of live and preserved specimens. (A) Male in resting colors, underwater photograph from
Guadalcanal, Solomon Islands; (B) flashing male in nuptial colors, underwater photograph from Nggatokae, western Solomon Islands; (C) flashing
male in nuptial colors, underwater photograph from the Solomon Islands; (D) AMS I.17479-001, 51.7 mm SL, male paratype, Tassafaronga Point,
Guadalcanal, Solomon Islands. Note purple spines and rays in preservation; (E) harem comprising one TP male (middle) and several females and
immature males, underwater photograph from Lovukol, central Solomon Islands. Photographs by M. Rosenstein (A–C, E) and Y. K. Tea (D).
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the Solomon Islands and recognized those from central
Indonesia (from Java east to Komodo and Sulawesi) as P.
paineorum, and those from southeastern South China Sea
(the Anambas Islands and Brunei) as P. x a n t h o c i r r i t u s .Thedis-
tinction of P. f i l a m e n t o s u s from P. p a i n e o r u m and P. x a n t h o c i r r i -
tus is supported by differences in TP male coloration pattern
and mitochondrial COI sequence data (Allen et al., 2016).
Based on the collection locality of the USNM 209923 para-
types, the identity of the specimens should be P. p a i n e o r u m ,
and not P. f i l a m e n t o s u s .
In addition to Papua New Guinea and the Solomon Islands,
Allen et al. (2016) reported on the occurrence of P. f i l a m e n t o s u s
in the far northern Great Barrier Reef of Australia. The record
appears to be spurious, as no photographs or examined mate-
rial from Australia were included in their study. The Australian
distribution record was also omitted from the distribution
map (fig. 18 in Allen et al., 2016). An extensive search of Aus-
tralian museum fish collections conducted by the first author
of the present study yielded only a single, previously unex-
amined specimen of P. filamentosus from Australia. The speci-
men was collected from Lizard Island, in the northern Great
Barrier Reef. The Lizard Island specimen (Fig. 10;AMS
I.40666-022) agrees well with P. filamentosus in meristic data,
morphometrics, dorsal- and caudal-fin morphology, and as-
pects of live and preserved coloration details (Fig. 11). The
Lizard Island specimen was compared with 12 paratypes and
four non-type specimens housed at the Australian Museum
(see list of examined material below). Unfortunately, the ho-
lotype (AMS I.16994-001) and four paratypes (AMS I.16995-
001) of P. filamentosus could not be located. An ongoing effort
to locate the missing types is currently being conducted by
the authors.
One record of P. filamentosus from the Gulf of Carpentaria
is listed on the Atlas of Living Australia, based on observa-
tions taken on the 2005 FRV Southern Surveyor expedition.
The specimen that the observation was based off could not
be located, and was likely unretained (J. Pogonoski, pers.
comm.). Unpublished data indicated that the specimen
weighed 19.5 g, which appears unlikely for a Paracheilinus.
We do not consider this record as verifiable. At this time, P.
filamentosus is known from Australia based solely on the
Lizard Island specimen. Given the specific habitat of Para-
cheilinus, it is likely that this species occurs more widely
across the northern Great Barrier Reef and the Coral Sea but
has evaded detection due to observation and sampling bias.
Indeed, the Lizard Island specimen was collected in Halimeda
fields growing in coral rubble on outer reef slopes, a habitat
not frequented by divers and fish collectors.
Etymology.—Named after the long filamentous segmented
dorsal-fin rays. Although this character is not diagnostic of
the species, it was at the time of its description. The only
other species of Paracheilinus known at the time was P. octo-
taenia, a species lacking filaments completely.
Paracheilinus flavianalis Kuiter and Allen, 1999
Yellowfin Flasher Wrasse
Figures 5C,8A–D,9
Paracheilinus sp. 1—Kuiter and Debelius, 1994: 232 (field
guide, underwater photograph from Indonesia).
Paracheilinus flavianalis Kuiter and Allen, 1999:123(holotype
WAM P.3 0 836-007, S c o t t R e e f, north-w e s t e r n s helf of
Wes tern A ustr ali a ); Kuiter and Tonozuka, 2004:496(field
guide, underwater photos from Indonesia); Michael, 2009:
287 (field guide, underwater photos from Indonesia);
Kuiter, 2010:115(fieldguide,underwaterphotosfrom
Indonesia); Allen and Erdmann, 2012:707(fieldguide,un-
derwater photo from Triton Bay, West Papua).
Diagnosis.—Dorsal-fin rays IX,11; anal-fin rays III,9; pecto-
ral-fin rays 14; pelvic-fin rays I,5; pored lateral-line scales
14–17 þ5–10 ¼19–27; gill rakers 4–6 þ7–10 ¼11–16; body
depth 3.1–3.5 in SL; head length 2.7–3.2 in SL; snout length
3.8–5.1 in HL; orbit diameter 3.0–4.6 in HL; interorbital
width 3.7–4.9 in HL; least depth of caudal peduncle 1.9–3.0
in HL; caudal peduncle length 1.6–2.5 in HL; TP males with
1–4 (usually one) elongate, red filamentous dorsal-fin rays,
longest dorsal-fin soft ray 1.8–3.6 in SL; pelvic fin length
1.9–2.9 in HL; caudal fin of TP males round without fila-
mentous lobes; caudal-fin length 3.8–4.4 in SL; body with
stripe pattern B; anal fin uniformly yellow or orangey yel-
low, basal third usually lined with a horizontal row of blue
spots (sometimes incomplete or completely absent).
Remarks.—Paracheilinus flavianalis is most similar to P.
amanda,P. carpenteri,P. mccoskeri, and P. rubricaudalis (see
comparisons in description of P. amanda above). The mem-
bers of this species complex are mostly allopatric, with P. fla-
vianalis overlapping only very narrowly with P. mccoskeri in
Bali and P. carpenteri in Sulawesi (Fig. 6C). It usually possesses
a single red dorsal-fin filament, but individuals may pos-
sess anywhere between one to four filaments (Fig. 8). Para-
cheilinus flavianalis is known from southern and eastern
Indonesia, from Bali eastwards across the Lesser Sunda Is-
lands, southern Sulawesi, the Moluccas, and West Papua. It
also occurs in Timor Leste. In Australia, it occurs in Evans
and Flinders Shoals, Timor Sea, off northeast Darwin,
Northern Territory, and Ashmore, Scott, Seringapatam, and
Hibernia Reefs in the north-western shelf of Western Aus-
tralia (Fig. 9).
Etymology.—The species is named flavianalis in reference to
the characteristic yellow anal fin.
Paracheilinus nursalim Allen and Erdmann, 2008
Nursalim’s Flasher Wrasse
Figures 12A–F
Paracheilinus nursalim Allen and Erdmann, 2008: 181 (holo-
type NCIP 6327, Triton Bay, West Papua, Indonesia);
Michael, 2009: 284 (field guide, underwater photograph
from Bird’s Head Peninsula, West Papua, Indonesia);
Kuiter, 2010: 109 (field guide, underwater photographs
from Raja Ampat and Triton Bay, West Papua, Indonesia);
Allen and Erdmann, 2012: 708 (field guide, underwater
photographs from Triton Bay, West Papua, Indonesia).
Diagnosis.—Dorsal-fin rays IX,11; anal-fin rays III,9; pecto-
ral-fin rays 14; pelvic-fin rays I,5; pored lateral-line scales 11–
16 þ3–10 ¼14–26; gill rakers 4–6 þ8–9 ¼12–15; body depth
2.9–3.6 in SL; head length 2.8–3.3 in SL; snout length 3.8–4.9
in HL; orbit diameter 3.0–4.2 in HL; interorbital width 3.9–5.3
in HL; least depth of caudal peduncle 2.0–2.5 in HL; caudal
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peduncle length 1.5–2.3 in HL; TP males with 3–6 (rarely 3)
elongate, yellow filamentous dorsal-fin rays (bright white
when in display), longest dorsal-fin soft ray 1.8–2.3 in SL; pel-
vic fin length 1.6–2.0 in HL; caudal fin of TP males with very
pronounced filamentous lobes, with filaments sometimes
greater than SL; caudal-fin length 1.6–2.4 in SL; caudal con-
cavity 0.8–1.5 in SL; body with stripe pattern A’; body with
two black rectangular epaulettes, one below spinous portion
of dorsal fin, one along lower caudal peduncle (dusky in pres-
ervation); ground color of body yellow orange in life; central
portion of caudal fin with intricate blue markings.
Remarks.—This beautiful species was previously known only
from the western Bird’s Head Peninsula of West Papua,
Indonesia, from Raja Ampat and Triton Bay (Fig. 12D–F). It
has also been photographed in Ambon, approximately 500
km west of the Bird’s Head Peninsula (Allen et al., 2016).
The Ambon record was based only on a single photograph,
and so it wasn’t clear if a viable population existed outside
of the Bird’s Head Peninsula. The discovery of P. nursalim in
the Timor Sea in northeastern Australia extends the known
distribution of this species approximately 1,000 km south of
Ambon, and approximately 1,200 km south of the Bird’s
Fig. 12. Paracheilinus nursalim, images of live and preserved specimens from Flinders Shoal, Timor Sea, off Darwin, Northern Territory, Australia
(A–C) and West Papua, Indonesia (D–F). (A) Flashing male in nuptial colors, aquarium specimen. Specimen not retained. (B) Male in resting colors,
aquarium specimen. Specimen not retained. (C) Male in preservation, NTM S.18257-001, 53.1 mm SL, Flinders Shoal, Timor Sea. (D) Flashing
male in nuptial colors, underwater photograph from Triton Bay, West Papua, Indonesia. (E) Male in resting colors, underwater photograph from
Triton Bay, West Papua, Indonesia. (F) AMS I.44190-001, 46.0 mm SL, male paratype, FakFak Peninsula, West Papua, Indonesia. Photographs by K.
Kohen (A); Y. K. Tea (B–C; F); E. Cheng (D); and L. Cheng (E).
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Head Peninsula. It is likely that the species occurs through-
out this region but has escaped detection until recently.
Australian records of P. n u r s a l i m are based on specimens
from Flinders Shoal, Timor Sea (Fig. 12A–C). Comparison of
live photographs and a single retained specimen (Fig. 12C;
NTM S.18257-001) agrees well with topotypical examples of P.
nursalim on the basis of meristic data, morphometrics, dorsal-
and caudal-fin morphology, and aspects of live and preserved
coloration details. Comparisons were made with two para-
types of P. n u r s a l i m (Fig. 12F; AMS I.44190-001). Some of the
Australian specimens appear to have the dorso-anterior black
epaulette below the spinous portion of the dorsal fin reduced
or absent completely, which is otherwise typical of this species
(Fig. 12A–B). However, this pattern appears to vary even in
specimens from Triton Bay (Fig. 12E). In all other aspects, they
are inseparable from topotypical examples of P. n u r s a l i m .We
attribute these differences in coloration to intraspecific varia-
tion, which is not uncommon for Paracheilinus.
Etymology.—Named nursalim at the request of Cherie Nursa-
lim and Michelle Liem, who successfully bid to support the
conservation of this species at the Blue Auction in Monaco
on 20 September 2007, and who have given generously to
support Conservation International’s Bird’s Head Seascape
marine conservation initiative. The name honors their par-
ents Sjamsul and Itjih Nursalim. The name nursalim is treat-
ed as a noun in apposition.
DISCUSSION
Four species of Paracheilinus are here reported as extant in
Australia, all of which confined to or around tropical lati-
tudes of Western Australia, Queensland, and the Northern
Territory (Fig. 6C). Despite their widespread occurrences in
these regions, representation of Australian species of Parachei-
linus is poor in Australian fish collections. Only P. flavianalis
is well represented in Australian collections and is known
from the holotype (WAM P.30836-007) and 17 paratypes de-
posited in WAM (WAM P.31194-001 [14]) and NMV (NMV
A20059 [3; erroneously reported as 2 specimens in Kuiter and
Allen, 1999]). It is also known from additional material col-
lected outside of Australia (see list of material examined).
Paracheilinus amanda is known primarily from the type se-
ries—the holotype at QM (QM I.39758) and five paratypes at
AMS (AMS I.50116-001 [1]), WAM (WAM P.33973.001 [2]), and
ZRC (ZRC 64175 [2]). Two additional non-type specimens are
also deposited in WAM (WAM P.34513.001) and ZRC (ZRC
64174). We are aware of one specimen of P. a m a n d a (incorrectly
identified as P. r u b r i c a u d a l i s )currentlydepositedintheIndone-
sian Biodiversity Research Centre under the voucher number
MB043301. The specimen was collected with the WAM para-
types from Harrier Reef, Great Barrier Reef, and not the Ribbon
Reefs as reported in Allen et al. (2016).Thespecimenwasnot
examined in this study, but mitochondrial COI sequence ob-
tained from this specimen was used in our phylogenetic analy-
ses and agrees well with sequences obtained from the AMS
paratype of P. a m a n d a (AMS I.50116-001). The species is also
known from an underwater photograph taken at Osprey Reef
in the Coral Sea (Fig. 5A2).
Paracheilinus filamentosus is known from Australia based
on a single specimen collected from Lizard Island, Great Bar-
rier Reef, currently deposited at AMS (AMS I.40666-022).
Twelve paratypes (AMS I.17479-001 [1] and AMS I.17496-
001 [11]) of P. filamentosus from the Solomons Islands are
deposited at AMS. The holotype (AMS I.16994-001) from
Papua New Guinea and four paratypes (AMS I.16995-001)
from the Solomon Islands could not be located.
Although Paracheilinus nursalim is known from numerous
photographs, only a single specimen is deposited in Austra-
lian fish collections. The specimen is a male collected from
Flinders Shoal, Timor Sea, currently deposited at NTM
(NTM S.18257-001).
Of the remote Australian Territories, Paracheilinus has been
reported only from the territories of Ashmore and Cartier Is-
lands (P. f l a v i a n a l i s [Allen, 1996;Kuiter and Allen, 1999]), and
the Coral Sea (P. a m a n d a ,presentstudy).Curiously,thegenus
has not been reported from the Christmas and Cocos (Keeling)
Islands despite a history of intensive surveying (Allen, 2000;
Allen et al., 2007;Hobbs et al., 2009,2010,2014;Hobbs and
Allen, 2014), especially given the neighboring presence of P.
mccoskeri and P. f l a v i a n a l i s in Bali, and P. f l a v i a n a l i s in the Less-
er Sunda Islands and the Timor Sea. It is likely that the absence
of Paracheilinus from this region, and other tropical regions of
Australia, is a result of sampling bias, as most species tend to
frequent somewhat deep rubble slopes in soft sediment habi-
tats away from coral reefs.
Alotof42unidentifiedspecimensofParacheilinus collected
from Tijou Reef, east coast of the Cape York Peninsula, Far
North Queensland is deposited at AMS (AMS I.20779-049).
However, these specimens could not be located, and as such
were unavailable for study and their identities could not be
verified.
MATERIAL EXAMINED
Paracheilinus amanda: QM I.39758, holotype, 47.6 mm SL,
Harrier Reef, Great Barrier Reef; AMS I.50116-001, paratype,
44.4 mm SL, Flora Reef, Coral Sea; WAM P.33973.001, 2, para-
types, 37.3–43.7 mm SL, Harrier Reef, Great Barrier Reef;
WAM P.34513.001, 54.3 mm SL, Harrier Reef, Great Barrier
Reef; ZRC 64174, 47.2 mm SL, off Taurama, southern Papua
New Guinea, Coral Sea; ZRC 64175, 2, paratypes, 32.3–47.6
mm SL, off Hula, southern Papua New Guinea, Coral Sea.
Paracheilinus carpenteri: BPBM 22424, holotype, 39.0 mm SL,
Mactan Island, Philippines; BPBM 21165, 2, paratypes,
35.0–36.8 mm SL, Buyong Beach, Mactan Island, Philip-
pines; BPBM 22116, paratype, 52.0 mm SL, Mactan Island,
Philippines; BPBM 22465, 10, paratypes, 21.6–65.8 mm SL,
Caban Island, Batangas, Philippines; BPBM 26507, 73.0 mm SL,
Sumilon Island, Philippines.
Paracheilinus filamentosus: AMS I.17479-001, paratype, 51.7
mm SL, Honiara, Guadalcanal, Solomon Islands; AMS
I.17481-013, 3, 23.0–37.1 mm SL (specimens cleared and
stained), near Tassafaronga Point, Guadalcanal, Solomon Is-
lands; AMS I.17496-001, 11, paratypes, 28.5–54.5 mm SL,
Alite Reef, off Malaita, Solomon Islands; AMS I.17521-004,
45.0 mm SL, Honiara, Guadalcanal, Solomon Islands; AMS
I.17524-004, 58.9 mm SL, Honiara, Guadalcanal, Solomon
Islands; AMS I.17530-001, 40.2 mm SL, Alite Reef, off
Malaita, Solomon Islands; AMS I.40666-022, 46.6 mm SL,
Lizard Island, Great Barrier Reef, Queensland, Australia.
Paracheilinus flavianalis:NMVA20059,3(erroneouslyreported
as 2 in Kuiter and Allen, 1999), paratypes, 50.4–51.2 mm SL,
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aquarium specimens from Bali; AMS I. 46121-035, 66.2 mm
SL, Christo Rei Beach, east of Dili, Timor Leste; AMS I.46121-
041, 40.4 mm SL, Christo Rei Beach, east of Dili, Timor Leste;
NTM S.17978-013, 35.9 mm SL, Evans Shoal, Timor Sea; NTM
S.18121-011, 54.3 mm SL, Evans Shoal, Timor Sea; NTM
S.18257-005, 31.7 mm SL, Flinders Shoal, Timor Sea; WAM
P. 3 1 1 9 4 . 0 0 1 , 5 ( o f 1 4 ) , 2 5 . 0 – 3 4 . 9 m m S L , c e n t r a l l a g o o n a t S e r -
ingapatam Reef, north-western Western Australia.
Paracheilinus lineopunctatus: AMS I.45300.194, 51.0 mm SL
(specimen cleared and stained), aquarium specimen from
the Philippines.
Paracheilinus mccoskeri: AMS I.18604-001, paratype, 37.8
mm SL, Moroni Reef, Ngazidja (Grand Comore), Comoros.
Paracheilinus nursalim: AMS I.44190-001, 2, paratypes, 40.6–
46 mm SL, Fak Fak Peninsula, Pulau Panjang, Barat Prov-
ince, West Papua, Indonesia; NTM S.18257-001, 53.1 mm
SL, Flinders Shoal, Timor Sea.
Paracheilinus paineorum: AMS I.34501-029, 21.0–42.0 mm
SL, Flores, Indonesia.
Paracheilinus rubricaudalis: WAM P.33979.001, 3, 40.8–52.7
mm SL, Pityilu Island, Manus Province, northern Papua New
Guinea; WAM P.34665.001, 2, 41.8–43.9 mm SL, Jacquinot
Bay, New Britain, Papua New Guinea.
ADDENDUM
At the time of writing, photographs of a species of Paracheili-
nus collected from American Samoa were sent to the senior
author by B. D. Greene and R. L. Pyle. The specimens closely
resemble P. rubricaudalis in coloration, but the genus (in-
cluding P. rubricaudalis) is not known from this locality.
These specimens were not available for examination at the
time of writing, and so their identities could not be deter-
mined. They will be dealt with in a forthcoming manuscript.
DATA ACCESSIBILITY
The tree file and the molecular sequence alignment used to
generate the maximum likelihood tree in this study are
available at: https://www.ichthyologyandherpetology.org/
i2023019.GenBankandBOLDaccessionnumbersarein-
cluded in the sequence alignment file. Unless an alternative
copyright or statement noting that a figure is reprinted
from a previous source is noted in a figure caption, the pub-
lished images and illustrations in this article are licensed by
the American Society of Ichthyologists and Herpetologists
for use if the use includes a citation to the original source
(American Society of Ichthyologists and Herpetologists, the
DOI of the Ichthyology & Herpetology article, and any individ-
ual image credits listed in the figure caption) in accordance
with the Creative Commons Attribution CC BY License.
ZooBank publication urn:lsid:zoobank.org:pub:FFB033F3-
092C-4654-92D4-8D4CA64BD832.
ACKNOWLEDGMENTS
We th a nk A. Ha y, I . Rile y, K . Park i nson , a nd S. Re a der (A M S), C.
Atta (BPBM), D. Bray (NMV), M. Hammer (NTM), J. Johnson
(QM), G. Moore and J. Ritchie (WAM), and K. Lim and H. H.
Tan (ZRC) fo r curat ori al as sista nce , loan of spec ime ns, and
provision of registration numbers, photographs, and X-rays.
Type specimens, as well as habitat information and distribution
records of the new species, were provided by T. Bennett, B.
Shutman, and Cairns Marine. Photographs used in this
manuscript were kindly provided by G. R. Allen, T. Cameron, J.
Castellano, V. Chalias, E. Cheng, A. DeLoach, N. DeLoach, K.
Endoh, T. Kawamoto, K. Kohen, R. H. Kuiter, J. Leis, T.
Miskiewicz, W. Osborn, M. Rosenstein, R. Smith, H. H. Tan,
and T. Yamazumi. A. C. Gill provided helpful comments on
interpretation of X-rays and osteology. This research was
supported by the Chadwick Biodiversity Research Fellowship,
awarded to Yi-Kai Tea in 2022. Established in 2008 with
thanks to a generous bequest from the late Clarence E.
Chadwick, the Chadwick Biodiversity Research Fellowship
provides opportunities for gifted young scientists to establish
a career in biodiversity research.
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