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Two New Species of Pseudojuloides from Western Australia and Southern
Japan, with a Redescription of Pseudojuloides elongatus (Teleostei: Labridae)
Yi-Kai Tea
1,2
, Anthony C. Gill
2,3
, and Hiroshi Senou
4
The anti-equatorial labrid Pseudojuloides elongatus has a wide but disjunct distribution across the Western Pacific and
Eastern Indian Oceans, with populations occurring in Western Australia, southern Japan, and the southwest Pacific
Ocean. Principal component analysis of morphological characters and coalescent-based species-tree estimates of
mitochondrial and nuclear DNA markers suggest that these populations are under incipient stages of divergence. The
three allopatric populations differ strongly in coloration patterns of both sexes, particularly in terminal males,
suggestive of reproductive isolation. We redescribe Pseudojuloides elongatus on the basis of nine paratypes and two
additional specimens from eastern Australia and Norfolk Island, and describe two new species, Pseudojuloides crux, new
species, from Western Australia, and P. paradiseus, new species, from southern Japan. The complex is distinguished from
other members of the genus in sharing the following combination of characters: body elongate; dorsal-fin rays IX,12;
pectoral-fin rays 12; no median predorsal scales; and usually 27 lateral-line scales. We briefly comment on anti-
equatorial biogeographical patterns and Pseudojuloides argyreogaster from the Western Indian Ocean.
FOWLER (1949) erected Pseudojuloides for Pseudojulis
cerasina Snyder (1904) on the basis of the following
characters that differ from Pseudojulis Bleeker (cur-
rently considered a synonym of Halichoeres [R¨
uppell,
1835]): scales on chest, breast, and space before pectoral
fins larger; shorter pectoral fins; caudal fin broadly scaled to
basal one-third; and coloration without a black lateral band
or spot at origin of the pectoral fin. The genus was later
expanded by Ayling and Russell (1977) with the description
of a second species, Pseudojuloides elongatus,fromNew
Zealand, Australia, and southern Japan. Randall and
Randall (1981) revised the genus, recognizing eight species,
including an additional five as new: Pseudojuloides atavai,P.
erythrops, P. mesostigma,P. p y r i u s ,andP. xanthomos. Since
then, eight additional new species have been described
from tropical Indo-Pacific reefs: P. e d w a r d i , P. k a l e i d o s ,P.
labyrinthus,P. p o l a c k o r u m ,P. p o l y n e s i c a ,P. severnsi,P.
splendens,andP. z eu s , bringing the total number of valid
species to 16.
Members of the genus are typically associated with coral
reefs in warm tropical waters. The exception is found in the
subtropical species Pseudojuloides elongatus, which inhabits
subtropical to temperate reefs dominated by seagrasses and
kelp (Randall and Randall, 1981). The species is represented
by three disjunct populations occurring in the Western
Pacific and eastern Indian Oceans: southern Japan, the
southwest Pacific Ocean, and Western Australia. This
distribution pattern agrees well with Randall’s (1981)
definition of anti-equatorial biogeographical patterns in
which species occupy geographical regions within tropical
latitudes, but not within equatorial waters. Often these taxa
are restricted to climatically cooler regions or have a
proclivity for cooler waters such as in thermoclines or
upwellings (Randall, 1981).
Some nominal species with widespread distributions
exhibit considerable variation in color patterns through-
out their range, raising the question of whether they are
truly widespread or are a complex of morphologically
similar cryptic species (Gill and Kemp, 2002; Tea et al.,
2019a; Tea and Gill, 2020). Closer evaluation of these
species has often revealed hidden diversity, leading to the
recognition of allopatric species (Victor, 2017; Walsh et al.,
2017; Tea and Gill, 2020). In the species description of
Pseudojuloides elongatus, Ayling and Russell (1977) included
specimens from New Zealand (type locality), eastern
Australia, Western Australia, and southern Japan. Kuiter
(2010) recognized that the Western Australian and Japa-
nese populations differed markedly from the nominal P.
elongatus in color patterns. However, no systematic work
has been done to address the taxonomic status of these
groups.
We herein revise the taxonomy of Pseudojuloides elongatus
on the basis of molecular and morphological data. Our
analyses indicate that the populations from Western Austral-
ia, southern Japan, and the southwest Pacific Ocean
represent a cryptic species complex in the early stages of
divergence. The strong geographical disjunction between
populations and consistent coloration differences in both
sexes suggest that these populations are reproductively
isolated. These patterns are similar to those in other species
of sexually dimorphic labrids where low levels of genetic
divergence accompany large differences in coloration and
other phenotypic characters (Victor and Randall, 2014).
Accordingly, we describe the populations from Western
Australia and southern Japan as new species. We also provide
brief comments on similar anti-equatorial biogeographical
patterns of distribution in other distantly related groups of
fishes and discuss the putative relationship of the lesser-
known Pseudojuloides argyreogaster.
1
School of Life and Environmental Sciences, University of Sydney, Sydney, Australia; Email: (YKT) yi-kai.tea@sydney.edu.au. Send reprint
requests to YKT.
2
Ichthyology, Australian Museum Research Institute, 1 William Street, Sydney, New South Wales 2010, Australia.
3
Chau Chak Wing Museum, Macleay Collections and School of Life and Environmental Sciences, A12—Macleay Building, University of
Sydney, New South Wales 2006, Australia; Email: anthony.c.gill@sydney.edu.au.
4
Kanagawa Prefectural Museum of Natural History, 499 Iryuda, Odawara, Kanagawa 250-0031, Japan; Email: senou@nh.kanagawa-museum.jp.
Submitted: 29 October 2019. Accepted: 9 April 2020. Associate Editor: M. T. Craig.
Ó2020 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-19-316 Published online: 9 October 2020
Copeia 108, No. 3, 2020, 551–569
MATERIALS AND METHODS
Morphological data.—Methods for counting and measuring
follow Victor and Randall (2014), with the following
modifications. Counts of principal caudal-fin rays follow Gill
et al. (2016): the uppermost principal caudal-fin ray is the ray
articulating with hypural 5, and the lowermost principal
caudal-fin ray is the ray articulating between the distal tips of
the parhypural and the haemal spine of preural centrum 2.
Counts of principal and branched caudal-fin rays are
presented as upper þlower, where the upper rays are those
associated with hypurals 3–5, and lower rays are those
associated with hypurals 1–2 and the parhypural. Procurrent
caudal-fin rays are those anterior (dorsal and ventral) to the
principal rays. Scales above the lateral line to the dorsal-fin
origin were counted posterodorsally from the first lateral-line
scale. Scales above the anal-fin origin to the lateral line were
counted anterodorsally from the anal-fin origin, and includ-
ed the small, often partially concealed scale near the fin
origin. Gill-raker counts were of the total number of outer
rakers on the first arch, including rudiments.
Osteological details were determined from X-radiographs.
Vertebral counts are presented as precaudal þcaudal. The
anteriormost vertebra with a haemal spine was considered as
the first caudal vertebra, the urostylar complex as the last.
Terminology of intermuscular bones follows Patterson and
Johnson (1995) and Johnson and Patterson (2001).
In the description for the new species, data are presented
for the holotype, followed by data for the paratypes in
parentheses. 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. Specimens are
deposited in the Australian Museum, Sydney (AMS),
Kanagawa Prefectural Museum of Natural History, Yokoha-
ma (KPM-NI), and Western Australian Museum, Perth
(WAM).
Meristic and morphometric data were analyzed in a
combined dataset using principal component analysis
(PCA) in JMP v14 (SAS Institute Inc., NC) from a total of
34 specimens (southwest Pacific Ocean, n¼11; southern
Japan, n¼10; Western Australia, n¼13). A total of 13
meristic and 25 morphometric characters were included. To
account for allometric and size differences between indi-
viduals, PCA of morphometric characters was based on
residual values obtained from pooled regression of each log
(ln)-transformed character against lnSL. Regression was
performed in RStudio v1.1.453 (RStudio Team, 2015). Log
transformation was used to reduce correlation between
standard length and variance. Missing morphometric values
were replaced with zero residual values. Scores of the most
informative principal components (PC I and PC II) were
visualized using bivariate plots. We further tested the
distinctiveness of each population by inputting eigenvec-
tors that explained the greatest percent variance from the
PCA into a multivariate analysis of variance (MANOVA)
implemented in the software PAST v.2.17 (Hammer et al.,
2001).
We made use of photographs from the Image Database of
Fishes housed at KPM-NI, which are assigned unique
numbers with the prefix KPM-NR. Note that owing to zero-
padding, a seven-digit number is used for the catalog number
in the fish specimen and the fish image collections of the
museum, but zero suppression is adopted for expression of
the essential numbers here.
Taxon sampling, DNA sequencing, and phylogenetic analyses.—
Tissue samples of Pseudojuloides elongatus from the southwest
Pacific Ocean (n¼1), Western Australia (n¼1), and southern
Japan (n¼1) were obtained from the right pelvic fin or caudal
fin of archived specimens, preserved in 100% ethanol, and
stored at –208C prior to DNA extraction. DNA was extracted
using the DNeasy Blood and Tissue kit (Qiagen) following the
manufacturer’s protocol. Three mitochondrial (cytochrome c
oxidase subunit 1 [COI], 16S ribosomal RNA [16S], and
control region) and two nuclear markers (ribosomal protein
S7 [S7] and recombination activating gene 2 [RAG2]) were
amplified using the polymerase chain reaction (PCR). The
extensive use of both nuclear and mitochondrial markers
allowed us to maximize the amount of information retriev-
able on account of the small number of archival museum
samples used. Primer sets and PCR conditions follow Chow
and Hazama (1998), Ahyong and Jarman (2009), Cheng et al.
(2012), DiBattista et al. (2012), and Chang et al. (2017) and
are presented in Supplemental Appendix A (see Data
Accessibility). Sanger sequencing of PCR products was out-
sourced to Macrogen (Seoul, South Korea). Forward and
reverse sequences were aligned and trimmed separately as
contigs using Geneious Prime 2019.1.1 (Biomatters). Gen-
Bank accession numbers are presented in Supplemental
Appendix B (see Data Accessibility).
We performed phylogenetic analyses of a dataset including
the aforementioned Pseudojuloides elongatus from the south-
west Pacific Ocean, Western Australia, and southern Japan,
with Halichoeres garnoti and Pseudojuloides atavai as out-
groups. Owing to the use of five genetic markers and the lack
of comparative sequence data for other species of Pseudoju-
loides, phylogenetic analyses were performed in order to
explore the taxonomic status of the new species, and do not
attempt to reconstruct the relationships among other species
within the genus. To account for potential gene-tree
incongruence due to incomplete lineage sorting common
among labrid species (Victor and Randall, 2014), we used the
program ASTRAL (Mirarab et al., 2014), which employs a
multi-species coalescent approach to estimate the species tree
given a set of unrooted gene trees. The ASTRAL species tree
was estimated using two gene trees corresponding to the 602
base pairs of nuclear S7 and to the 2,027 base pairs of COI,
16S, and control region concatenated into a single mito-
chondrial alignment. Sequences were aligned using the
MUSCLE v3.8.31 algorithm (Edgar, 2004), and unrooted
gene trees were inferred using maximum-likelihood analyses
conducted in RAxML v8.2.12 (Stamatakis, 2014). The gene
tree for the nuclear RAG2 marker was not considered, because
sequences were only successfully amplified for P. elongatus
from Western Australia and southern Japan. These sequences
were compared in a pairwise fashion instead (p-distance). In
addition to ASTRAL, we analyzed the concatenated S7 and
mitochondrial dataset using maximum-likelihood conducted
in RAxML. The analysis was performed in duplicate to check
for local optima, each using ten random starts. Bootstrapping
was performed using 1,000 pseudoreplicates of the data. The
best-fitting substitution models and data-partitioning
scheme were selected using PartitionFinder2 (Lanfear et al.,
2016).
552 Copeia 108, No. 3, 2020
RESULTS
Principal component analysis and the multispecies coalescent.—
Principal component analysis of morphometric and meristic
characters revealed overlap between individuals from south-
ern Japan and the southwest Pacific Ocean (Fig. 1A), but a
clear separation in the Western Australian individuals on PC
II (P,0.0001, MANOVA on first four axes explaining 55.2%
of total variance). The greatest discrimination was offered by
PC I (23.5% of variance; Fig. 1A; Table 1), with individuals
from Western Australia and southern Japan on average
having more positive scores than those from the southwest
Pacific Ocean (Fig. 1B). The characters with the highest
loadings along PC I were lengths of first and second dorsal-
fin spines, third anal-fin spine, and caudal-fin length (Fig. 1B;
Table 1). Explaining 16% of variance, PC II primarily indexed
variation in head length, predorsal length, and snout length.
Individuals from Western Australia were more clearly
separated from the other populations in having more
negative scores on PC II.
Comparison of mitochondrial DNA revealed the greatest
differences in the control region, with a 2.1% difference (p-
distance) between the individuals from Western Australia
and southern Japan, and between the individuals from
southern Japan and the southwest Pacific Ocean. No
nucleotide differences in the control region were detected
between the individuals from Western Australia and the
southwest Pacific Ocean. Pairwise distances for all remaining
mitochondrial markers were less than 1%, with the largest
difference observed between the individuals from Western
Australia and the southwest Pacific Ocean (p-distance ¼0.9%
at COI; Appendix 1). Comparison of nuclear DNA for all
sampled individuals revealed the greatest difference in S7,
with a 1.3% difference between the individuals from Western
Australia and the southwest Pacific Ocean (Appendix 2).
Because we were unable to retrieve sequences of P. elongatus
from the southwest Pacific Ocean, a pairwise comparison of
the nuclear RAG2 marker was performed only for individuals
from Western Australia and southern Japan (p-distance ¼
1.3% at RAG2).
We used a multispecies-coalescent approach in ASTRAL to
infer the species tree given the unrooted gene trees from the
nuclear S7 gene tree and the concatenated mitochondrial
COI,16S, and control region. The species tree supported a
sister relationship between individuals from the southwest
Pacific and southern Japan, consistent with the results from
the PCA of morphometric and meristic characters (Fig. 1C).
Pseudojuloides elongatus Ayling and Russell, 1977
Long Green Wrasse
Figures 2A, 3, 4; Table 2
Pseudojuloides elongatus Ayling and Russell, 1977: 174, figs 4–
6, 8–10 (holotype NMNZ 6153, Poor Knights Islands, New
Zealand; paratypes from New Zealand, Norfolk Island, and
New South Wales, but not those from Japan or Western
Australia); Randall and Randall, 1981: 56 (revision);
Francis, 1993: 165 (checklist); Parenti and Randall, 2000:
36 (checklist); Fricke et al., 2011: 418 (New Caledonia);
Russell, 2015: 1386, fig. 202.15 (description, southwest
Pacific distribution only; color photos); Kuiter and Kuiter,
2018: 250 (distribution, color photos).
Diagnosis.—A species of Pseudojuloides, distinguished primar-
ily from all other species in the genus based on the following
combination of characters: no median predorsal scales; a
large canine tooth at corner of mouth; body of males
primarily olive-green, upper body with 0–4 rows of metallic
blue spots, not overlain in black; females unmarked,
olivaceous overall.
Description.—Dorsal-fin rays IX,12–13 (usually IX,12), all
segmented rays branched; anal-fin rays III,12, all segmented
rays branched; pectoral-fin rays 12, upper two rays and
sometimes lower one ray unbranched; pelvic-fin rays I,5, all
Fig. 1. (A) Principal component analysis of 13 meristic and 25 morphometric characters and (B) loading plot for all specimens of Pseudojuloides
elongatus (sensu lato) from Western Australia (n= 13), southwest Pacific Ocean (n= 11), and southern Japan (n= 10). AO = anal-fin origin; DO =
dorsal-fin origin; LL = lateral line. (C) Species tree estimated using a multispecies-coalescent approach in ASTRAL. Branch lengths are estimated only
for internal branches (in coalescent units), which directly measure the amount of discordance between gene trees. Lengths of terminal branches and
the internal branch descending from the root are arbitrary. Values at nodes correspond to local posterior probabilities and likelihood bootstrap
support, respectively.
Tea et al.—Revision of the Pseudojuloides elongatus complex 553
segmented rays branched; principal caudal-fin rays 7þ7,
uppermost and lowermost unbranched; upper procurrent
caudal-fin rays 5–6; lower procurrent caudal-fin rays 5–6;
total caudal-fin rays 24–26; lateral-line scales 26–27 (usually
27), posteriormost overlying hypural joint, with an addition-
al large tubed scale on caudal fin; scales above lateral line to
origin of dorsal fin 3–4; scales above anal-fin origin to lateral
line 8–10; circumpeduncular scales 16; gill rakers 14–17;
branchiostegals 6.
Vertebrae 9þ16; first vertebral centrum about half width of
subsequent centra, bearing a relatively short neural spine;
single supraneural, inserted in first interneural space; first
dorsal ptergiophore bearing one supernumerary spine and
one serial spine, inserted along with second pterygiophore in
second interneural space; remaining dorsal pterygiophores
insert one per interneural space, except for 18
th
interneural
space with two pterygiophores; first two haemal spines with
enlarged secondary haemal arches (see Russell, 1988: fig. 23),
first recurved towards second haemal spine; first anal
pterygiophore bearing two supernumerary spines and one
serial spine, inserted anterior to first haemal spine; subse-
quent anal pterygiophores each bear a serially associated
segmented ray and inserted one per interhaemal space,
except for two pterygiophores in 11
th
interhaemal space; ribs
present on vertebrae 3 through 9; epineurals present on
vertebrae 1 through 13–14; preural 2 haemal spine autoge-
nous; hypurals 1 and 2 undifferentiated from each other;
hypurals 3 and 4 undifferentiated from each and from
urostylar complex; hypural 5 autogenous, with broad keel-
like projection anteriorly; parhypural autogenous without
apparent hypurapophysis (Fig. 2A).
Body elongate, greatest depth 4.3–5.3 in standard length
(SL), width 1.7–2.2 in greatest depth; head pointed, length
3.1–3.5 in SL; dorsal profile of head slightly convex; snout
moderately long, 2.7–3.4 in head length (HL); caudal
peduncle short and narrow, least depth 2.6–3.1 in HL, its
length 2.7–3.5 in HL. Mouth small, terminal, upper jaw
nearly reaching a point vertically below anterior nostril. A
pair of large, moderately projecting, and slightly recurved
canines present anteriorly in each jaw, upper pair slightly
flared and widened, lower pair curving forward and fitting
between upper canines when mouth closed; a short row of 2–
7 (upper jaw) and 4–9 (lower jaw) irregularly placed chisel-
like incisiform teeth along each side of jaws; a canine tooth
sometimes present posteriorly on upper jaw of large
specimens.
Preopercular margin smooth, upper part free to between
level of corner of mouth and lower orbit; lower margin free
anterior to, or anterior to front edge of, orbit. Nostrils small,
anterior to upper edge of orbit; anterior nostril in a short
membranous tube, posterior nostril nearly covered by a
dermal flap from anterior edge. Lateral line continuous,
angling sharply downward beneath soft portion of dorsal fin
to straight peduncular portion; lateral-line scales with a
single pore.
Scales cycloid, moderately large except those on thorax
and nape, which are much smaller; head naked; fins unscaled
except basal portion of caudal fin and base of pelvic fins.
Dorsal- and anal-fin spines progressively longer, last dorsal-
fin spine 3.1–3.8 in HL and 1.2–1.5 in longest soft dorsal-fin
ray, third anal-fin spine 3.5–5.0 in HL and 1.3–1.9 in longest
soft anal-fin ray; caudal fin truncate to slightly rounded in
females and males; caudal fin relatively small, 1.4–1.5 in HL;
pectoral fins small, 2.0–2.3 in HL; origin of pelvic fins below
lower base of pectorals, their length 2.2–2.6 in HL. Morpho-
metric values are summarized in Table 2.
Coloration of males in life.—Based on color photographs of
live individuals in the field and aquaria (Fig. 3). Upper head
and body green to olive; lower part of head yellow-green to
ochre; head with a network of metallic blue to lilac markings;
interorbital and upper part of snout with 3–4 metallic blue to
lilac stripes, often sinuous and reaching upper lip; a pair of
metallic blue to lilac stripes present from behind upper orbit
Table 1. Character loadings and percent variance explained for the first
four principal component axes for all morphometric and meristic
measurements of all specimens of Pseudojuloides elongatus examined
for this study, Western Australia (n¼13), southwest Pacific (n¼11),
and southern Japan (n¼10). Highest loadings are indicated in bold. AO
¼anal-fin origin; DO ¼dorsal-fin origin; LL ¼lateral line.
Morphometric and
meristic characters
PC loading values
Axis 1 Axis 2 Axis 3 Axis 4
Percent variance explained 23.50% 16.0% 7.88% 7.84%
Greatest body depth 0.291 0.510 –0.176 –0.112
Depth of dorsal-fin origin 0.315 0.615 0.054 0.294
Body width 0.211 –0.152 0.129 –0.390
Head length 0.264 0.834 0.317 0.081
Snout length 0.287 0.706 0.261 0.130
Orbit diameter 0.385 0.575 –0.025 –0.159
Interorbital width 0.262 0.449 0.161 0.075
Caudal peduncle length 0.290 –0.389 0.388 0.483
Caudal peduncle depth 0.216 0.225 –0.005 –0.372
Predorsal length 0.286 0.745 0.397 0.140
Preanal length 0.053 0.513 0.347 –0.327
Prepelvic length 0.137 0.664 0.353 –0.198
Dorsal-fin base 0.289 –0.707 –0.004 0.023
1
st
dorsal-fin spine 0.931 –0.009 –0.167 –0.110
2
nd
dorsal-fin spine 0.937 0.107 –0.079 –0.070
Longest dorsal-fin ray 0.342 0.204 0.027 0.710
Anal-fin base 0.181 –0.102 –0.632 0.373
1
st
anal-fin spine 0.523 0.279 –0.208 0.710
2
nd
anal-fin spine 0.823 –0.363 0.295 0.001
3
rd
anal-fin spine 0.942 –0.155 –0.018 –0.111
Longest anal-fin ray 0.923 –0.201 –0.022 –0.085
Caudal fin length 0.945 –0.169 0.052 –0.117
Pectoral fin length 0.897 –0.101 0.079 –0.105
Pelvic-fin spine 0.349 0.322 –0.397 0.470
Pelvic fin length 0.307 0.388 –0.247 0.650
Number of dorsal-fin spines 0.167 0.200 –0.189 –0.304
Number of dorsal-fin rays 0.321 –0.122 0.053 0.175
Number of anal-fin rays 0.072 –0.535 0.618 0.121
Number of pectoral-fin rays 0.075 –0.131 0.619 0.120
Upper procurrent rays –0.053 –0.094 0.619 0.480
Lower procurrent rays –0.137 –0.327 0.522 0.113
Number of lateral–line scales 0.121 0.090 0.224 –0.242
Number of scales from LL
to DO
0.552 –0.130 –0.186 0.017
Number of scales from LL
to AO
0.187 –0.306 0.313 0.371
Upper gill rakers 0.025 0.104 0.015 –0.289
Lower gill rakers 0.362 –0.336 –0.245 0.137
Teeth on upper jaw 0.702 –0.256 –0.164 –0.308
Teeth on lower jaw 0.598 –0.590 0.143 –0.032
554 Copeia 108, No. 3, 2020
to upper edge of operculum; a second pair of stripes of the
same color present obliquely from lower jaw and lower orbit
to lower part of operculum; upper part of nape and
sometimes narrow area before dorsal fin with additional lilac
stripe; iris bright yellow to orange, with blue submarginal
ring around pupil; pectoral-fin base with conspicuous yellow
to black band, edged distally in metallic blue; lower part of
abdomen yellowish green; dorsal half of body green to olive
with 0–4 rows of metallic pink to blue spots; spots often
increasingly broken and irregular ventrally; ventral half of
body green without any obvious markings; dorsal fin orange
to pinkish orange with anterior interspinous membrane
spaces green, breaking up into oblique bands toward
posterior three-quarters of fin; distal margin of fin narrowly
bright blue; anal fin similar to dorsal fin, but with irregular,
oblique lilac to light blue bands; distal margin of fin bright
blue; caudal fin translucent green, sometimes with metallic
blue spots near base; pelvic fins hyaline blue-gray, spines and
rays metallic blue, with two bright red spots present between
each intermembrane space; pectoral fins pinkish hyaline.
Coloration of females in life.—Based on color photographs of
live individuals in the field (Fig. 4). Upper head and body
green to olive; lower part of head yellow-green; head and
body without any markings; iris bright yellow to orange,
with blue submarginal ring around pupil; dorsal, anal, and
caudal fins unmarked hyaline green; pelvic fins hyaline;
pectoral fins translucent.
Coloration in alcohol.—Uniformly pale tan; pectoral fin axil
spot black; metallic blue spots on back dark tan; dorsal and
anal fins hyaline, markings fading to tan; pelvic fins hyaline,
red spots fading to dark brown.
Habitat and distribution.—Pseudojuloides elongatus is known
from the southwest Pacific Ocean. Along the eastern
Australian coast, the species occurs in New South Wales to
Montague Island. It also occurs in Lord Howe Island, Norfolk
Island, New Caledonia, and in northeastern New Zealand,
from Cape Reinga to Poor Knights Island (Fig. 5). It frequents
kelp and other weedy areas at depths ranging from 2–25 m.
Etymology.—The species was named elongatus in reference to
its elongate body form.
Material examined.—New South Wales, Australia: AMS
I.17033-036, 48.6 mm SL (paratype), Sydney Harbour, North
Head, 338490S, 1518170E, 7 m, G. R. Allen, D. F. Hoese, G.
McPherson, J. Paxton, D. Pollard, and B. C. Russell, 6 April
1974; AMS I.17735-008, 56.5 mm SL (paratype), Sydney
Harbour, Camp Cove, 338500S, 1518160E, 5 m, R. H. Kuiter, 28
April 1974; AMS I.17743-004, 3, 55.3–69.3 mm SL (para-
types), Sydney Harbour, Watsons Bay, 338500S, 1518160E, 5 m,
handnet, R. H. Kuiter, 13 April 1974; AMS I.17800-001, 3,
64.9–73.3 mm SL (paratypes), Seal Rocks, 328260S, 152832 0E,
8 m, R. H. Kuiter, handnet, 13 May 1974; WAM P.27074-003,
1, 87.5 mm SL, Byron Bay, north-west side of Julian Rocks,
288380S, 1538370E, J. B. Hutchins, spear, 21 December 1980;
WAM P.27076-017, 1, 121.8 mm SL, Byron Bay, Julian Rocks,
288380S, 1538370E, J. B. Hutchins, spear, 23 December 1980.
Norfolk Island, Australia: AMS I.18772-001, 64.3 mm SL
(paratype), north side of Phillip Island, 15 m, B. C. Russell
and A. Piper, 20 September 1975.
Fig. 2. X-radiographs of Pseudoju-
loides elongatus (sensu lato). Images
not to scale. (A) AMS I.17743-004,
69.3 mm SL, male paratype, Watsons
Bay, Sydney Harbor; (B) WAM
P.25368-001, 100.7 mm SL, Tanta-
biddy Creek, Western Australia; (C)
KPM-NI 43448, 103.0 mm SL, Izu
Oceanic Park. Radiographs by A. Hay
and S. E. Reader (A, B) and H. Senou
(C).
Tea et al.—Revision of the Pseudojuloides elongatus complex 555
Fig. 3. Underwater photographs of Pseudojuloides elongatus (sensu stricto) from eastern Australia: (A) male, approximately 120 mm TL, 8 m, North
Solitary Island, New South Wales, Australia; (B) male, approximately 150 mm TL, 10 m, Cabbage Tree Bay, Manly Beach; (C) male, approximately 100
mm TL, aquarium specimen collected from Sydney Harbor. Specimen not retained. Photographs by I. Shaw, J. Sear, and R. H. Kuiter, respectively.
556 Copeia 108, No. 3, 2020
Pseudojuloides crux, new species
urn:lsid:zoobank.org:act:18B36E8A-4809-4C8E-8D23-
1395C4F572D0
Stellate Pencil Wrasse
Figures 2B, 6, 7; Table 3
Pseudojuloides elongatus (non Ayling and Russell, 1977): Allen,
1985: 2512, figs. 352, 353 (checklist; color photos; North
West Cape, Western Australia); Parenti and Randall, 2000:
36 (checklist).
Pseudojuloides sp. 3: Kuiter, 2010: 338 (color photo; Shark Bay,
Western Australia, Australia).
Holotype.—(Fig. 6A) WAM P.25368-001, 100.7 mm SL, male,
Western Australia, North-West Cape, lagoon reef off Tanta-
biddy Creek, G. R. Allen, spear, 25 June 1975.
Paratypes.—(Figs. 6B, 7) AMS I.48996-001,97.7 mm SL,
Western Australia, Dampier Archipelago, southwest of Eagle-
hawk Island, J. Gill, hand net, 6 May 2018; WAM P.25110-
001 (also paratypes of P. elongatus), 3, 87.0–97.3 mm SL,
Western Australia, Dampier Archipelago, Kendrew Island,
20828.58S, 1168320E, G. R. Allen and R. C. Steene, rotenone
and spear, 2 November 1973; WAM P.25318-007 (also
paratype of P. elongatus), 104.9 mm SL, Western Australia,
Houtman Abrohlos, Wallabi Group, Batavia wreck site, 8–10
Fig. 4. Underwater photograph of Pseudojuloides elongatus (sensu stricto), female, approximately 100 mm TL, 9 m, North Solitary Island, New
South Wales, Australia. Photo by I. Shaw.
Fig. 5. Distribution records for Pseu-
dojuloides elongatus (sensu lato):
Western Australia (pink), southern
Japan (purple), and southwest Pa-
cific (blue). Type locality is repre-
sented by an outlined circle. The
equator and tropic lines are repre-
sented by solid and dashed orange
lines, respectively.
Tea et al.—Revision of the Pseudojuloides elongatus complex 557
Table 2. Morphometric values for Pseudojuloides elongatus expressed as percentage SL.
AMS
I.17033-036
(paratype)
AMS
I.17743-004
(paratype)
AMS
I.17735-008
(paratype)
AMS
I.17743-004
(paratype)
AMS
I.18772-001
(paratype)
AMS
I.17800-001
(paratype)
AMS
I.17743-004
(paratype)
AMS
I.17800-001
(paratype)
AMS
I.17800-001
(paratype)
WAM
P.27074-003
WAM
P.2706-017
Standard length (mm) 48.6 55.3 56.5 60.3 64.3 64.9 69.3 72.0 73.3 87.5 121.8
Greatest body depth 19.5 19.5 19.6 19.7 20.1 20.2 19.3 20.4 18.8 21.2 23.3
Body depth at dorsal
origin
17.9 16.6 17.5 17.9 18.5 18.6 17.5 18.8 17.6 20.0 21.3
Body width 10.5 9.9 10.4 10.8 9.8 11.1 11.1 10.4 9.7 11.0 10.7
Head length 31.1 29.7 31.0 31.8 31.9 30.7 28.9 29.6 30.0 31.1 30.3
Snout length 9.3 9.6 10.4 11.3 11.4 10.6 10.1 10.6 10.4 11.3 11.3
Orbit diameter 6.4 6.5 6.2 6.0 6.4 5.9 5.9 5.3 5.3 5.5 4.4
Interorbital width 6.2 6.0 6.0 6.1 5.9 6.0 5.6 6.0 6.3 5.7 6.2
Caudal-peduncle depth 10.7 10.7 10.6 10.3 10.6 10.6 11.0 9.4 9.8 11.5 11.2
Caudal-peduncle length 9.7 8.9 9.9 9.1 10.1 10.3 9.7 11.0 9.8 9.0 8.8
Predorsal length 29.8 27.8 29.9 29.5 30.9 30.2 28.4 29.3 27.8 29.8 29.4
Preanal length 51.0 52.8 51.7 53.1 52.4 52.1 50.5 49.9 49.7 52.9 51.8
Prepelvic length 32.1 31.8 32.3 32.3 33.0 32.2 30.4 32.2 31.7 32.6 33.3
Base of dorsal fin 59.1 57.7 56.5 55.1 57.4 56.8 57.6 56.5 57.0 57.4 59.6
First dorsal-fin spine 4.5 4.2 4.1 4.5 3.7 4.0 4.3 4.0 4.6 3.5 4.5
Second dorsal-fin spine 5.6 5.1 5.8 5.6 5.4 5.4 5.5 5.6 6.4 5.3 5.8
Last dorsal-fin spine 9.1 8.5 8.3 8.5 8.4 9.6 8.2 9.4 9.4 8.1 8.9
Longest dorsal-fin ray
(number)
11.7 (10) 11.9 (10) 11.7 (10) 11.6 (10) 12.4 (9) 12.6 (9) 11.8 (10) 12.9 (2) 12.3 (2) 11.9 (9) 12.0 (10)
Base of anal fin 37.7 38.0 37.9 37.6 39.0 38.2 38.2 40.4 40.0 37.0 40.4
First anal-fin spine 3.5 3.8 3.0 4.0 3.1 2.9 3.3 3.3 2.7 3.0 3.1
Second anal-fin spine 6.8 5.8 5.7 6.0 6.2 5.7 5.3 5.3 5.3 4.7 4.6
Third anal-fin spine 8.2 7.4 7.8 8.1 9.2 8.0 6.9 7.1 7.4 6.2 6.1
Longest anal-fin ray
(number)
11.5 (4) 11.8 (3) 10.4 (3) 11.3 (3) 11.8 12.0 (3) 10.8 (3) 12.4 (3) 12.8 (2) 9.6 (3) 11.7 (3)
Caudal-fin length 21.4 21.5 20.5 22.1 22.9 22.3 20.9 21.0 21.4 20.6 20.5
Pectoral-fin length 13.8 13.9 13.3 13.6 14.5 14.5 12.7 14.6 13.4 14.1 13.9
Pelvic-spine length 7.8 7.4 7.8 7.8 8.7 8.6 7.1 7.8 9.0 7.7 8.7
Pelvic-fin length 11.9 11.9 12.0 12.3 13.1 13.4 11.8 13.5 13.6 12.6 13.4
558 Copeia 108, No. 3, 2020
m, G. R. Allen, spear, 22 May 1975 (field number ABR-12);
WAM P.25318-008 (also paratype of P. elongatus), 111.4 mm
SL, collected with WAM P.25318-007; WAM P.25825-002,
74.1 mm SL, Western Australia, South Murion Island, J. B.
Hutchins, 12 June 1977 (field number MUR-77-015); WAM
P.25368-038, 3, 79.2–103.8 mm SL, collected with holotype;
WAM P.25826-040, 65.6 mm SL, Western Australia, South
Murion Island, J. B. Hutchins and J. Trendall, 12 June 1977
(field number MUR-77-016); WAM P.26649-004, 96.6 mm SL,
Western Australia, Shark Bay, South Passage, south-east of
Monkey Rock, 268090S, 1138100E, 3–4 m, J. B. Hutchins, spear,
4 April 1979.
Diagnosis.—Pseudojuloides crux belongs to the Pseudojuloides
elongatus complex, species of which differ primarily from all
other species of Pseudojuloides in lacking median predorsal
scales and in having a large canine tooth on corner of mouth.
It shares overlapping or nearly overlapping meristic counts
and morphometric proportions with Pseudojuloides elongatus,
but differ in having males that are greenish yellow to
yellowish orange, upper body strongly overlain in black,
with 3–5 rows of metallic blue spots, and unmarked females
that are pinkish orange to olivaceous.
Description.—For data and description that do not differ
between species of this complex, see description of Pseudo-
juloides elongatus. Dorsal rays IX,12, all segmented rays
branched; anal-fin rays III,12 (one paratype with III,13), all
segmented rays branched; pectoral-fin rays 12 (one paratype
with 11 on one side; one paratype with 13 on one side),
upper two unbranched; principal caudal-fin rays 7þ7; upper
and lowermost unbranched (one paratype with 7 branched
rays ventrally); upper procurrent caudal-fin rays 6 (three
paratypes with 5); lower procurrent caudal-fin rays 5 (three
paratypes with 6); total caudal-fin rays 25 (three paratypes
with 24; three paratypes with 26); lateral line with 27 tubed
scales (one paratype with 26 tubed scales on one side),
posteriormost overlying hypural joint, with an additional
large tubed scale on caudal fin; scales above lateral line to
origin of dorsal fin 4; scales above anal-fin origin to lateral
line 8/9 (10 on one side of one paratype, 9 in all other
paratypes); gill rakers 17 (15–18). Vertebrae 9þ16; epineurals
present on vertebrae 1 through 13 (1 through 11 in one of
three paratypes examined; Fig. 2B).
Body elongate, greatest depth 4.3 (4.3–5.2) in SL, width 2.3
(1.8–2.4) in greatest depth; head pointed, length 3.2 (3.0–3.4)
in SL; dorsal profile of head slightly convex; snout moder-
ately long, 2.7 (2.6–3.1) in HL; caudal peduncle short and
narrow, least depth 2.8 (2.7–3.1) in HL, its length 3.1 (2.9–
4.1) in HL. Mouth small, terminal, upper jaw nearly reaching
a point vertically below anterior nostril. A pair of large,
moderately projecting, and slightly recurved canines present
anteriorly in each jaw, upper pair slightly flared and widened,
lower pair curving forward and fitting between upper canines
when mouth closed; a short row of 5–8 (upper jaw) and 6–9
Fig. 6. Underwater photographs of Pseudojuloides crux, new species, from a reef off Tantabiddy Creek, North-West Cape, Western Australia: (A)
male holotype, WAM P.25368-001, 100.7 mm SL; (B) female paratype, WAM P.25368-038, 79.2 mm SL. Photographs by G. R. Allen.
Tea et al.—Revision of the Pseudojuloides elongatus complex 559
(lower jaw) irregularly placed chisel-like incisiform teeth
along each side of jaws; a canine tooth sometimes present
posteriorly on upper jaw of large specimens.
Scales cycloid, moderately large except those on thorax
and nape, which are much smaller; head naked; fins
unscaled except basal portion of caudal fin and base of
pelvic fins. Dorsal- and anal-fin spines progressively
longer, last dorsal-fin spine 2.9 (2.5–3.6) in HL and 1.3
(1.2–1.6) in longest soft dorsal-fin ray, third anal-fin spine
4.2 (2.5–5.1) in HL and 1.9 (1.6–1.9) in longest soft anal-fin
ray; caudal fin rounded, relatively small, 1.3 (1.3–1.5) in
HL; pectoral fins small, 2.2 (2.0–2.3) in HL; origin of pelvic
fins below lower base of pectorals, their length 2.3 (2.2–
2.5) in HL. Morphometric values are summarized in Table
3.
Coloration of males in life.—Based on color photographs of
specimens when freshly dead, and live individuals in the
field (Figs. 6A, 7A): upper head and body greenish yellow to
yellowish orange; lower part of head yellowish green; head
with a network of lilac to light blue markings, often metallic;
interorbital and upper part of snout with 3–4 lilac to light
blue stripes, often sinuous and reaching upper lip; a pair of
metallic blue to lilac stripes present from behind upper orbit
to upper edge of operculum; a second pair of stripes of the
same color present obliquely from lower jaw and lower orbit
to lower part of operculum; upper part of nape and
sometimes narrow area before dorsal fin with additional lilac
to light blue stripe; iris bright yellow to orange, with blue
submarginal ring around pupil; pectoral-fin base with
conspicuous yellow to black band, edged distally in metallic
blue; lower part of abdomen greenish yellow to yellowish
orange; dorsal half of body yellow to yellowish green,
overlain with black, and with 4–5 rows of metallic blue
spots; ventral half of body yellow to yellowish green without
any obvious markings; dorsal fin bright orange to pinkish
orange with anterior interspinous membrane spaces bright
green, breaking up into oblique bands toward posterior three-
quarters of fin; distal margin of fin narrowly metallic blue;
anal fin similar to dorsal fin, but with irregular, oblique lilac
to light blue bands; distal margin of fin metallic blue; caudal
fin translucent yellow to yellow-green, sometimes with
metallic blue spots present near base; pelvic fins hyaline
blue-gray, spines and rays light blue, with bright red spots
present between each intermembrane space; pectoral fins
yellowish hyaline.
Coloration of females in life.—Based on color photographs of
live individuals in the field (Fig. 6B): upper head and body
olive; lower part of head orangey green; head and body
without any markings; iris bright yellow to orange, with blue
submarginal ring around pupil; dorsal, anal, and caudal fins
unmarked hyaline green; pelvic fins hyaline; pectoral fins
translucent.
Coloration in alcohol.—(Fig. 7B) Freshly preserved specimens
with blue tinge, fading to tan over time; black markings
remain; pectoral fin axil spot becomes black; metallic blue
spots on back become pale tan; nape dusky; sinuous
Fig. 7. Paratype of Pseudojuloides crux, new species, AMS I.48996-001, 97.7 mm SL, male paratype collected from southwest of Eagle Hawk Island
in the Dampier Archipelago. (A) Freshly euthanized showing post-mortem coloration; (B) freshly preserved showing bluish coloration. Photographs
by J. Gill and Y. K. Tea, respectively.
560 Copeia 108, No. 3, 2020
Table 3. Morphometric values for Pseudojuloides crux expressed as percentage SL.
Holotype Paratypes
WAM
P.25368-
001
WAM
P.25826-
040
WAM
P.25825-
002
WAM
P.25368-
001
WAM
P.25110-
001
WAM
P.25110-
001
WAM
P.26649-
004
WAM
P.25110-
001
AMS
I.48996-
001
WAM
P.25368-
038
WAM
P.25368-
038
WAM
P.25318-
007
WAM
P.25318-
008
Standard length
(mm)
100.7 65.6 74.1 79.2 87.0 94.2 96.6 97.3 97.7 101.7 103.8 104.9 111.4
Greatest body
depth
23.1 19.1 20.5 21.1 22.9 22.9 20.8 20.8 22.9 22.1 21.1 21.1 21.8
Body depth at
dorsal origin
21.7 19.1 20.0 19.4 20.7 19.8 20.1 20.8 21.6 20.8 20.1 20.0 19.7
Body width 10.1 10.2 10.5 10.0 10.2 10.4 11.6 10.4 10.1 9.4 10.3 10.1 9.1
Head length 30.9 32.5 32.5 31.3 33.0 32.5 30.8 32.4 31.7 31.3 29.7 29.9 29.9
Snout length 11.6 11.4 11.3 11.6 11.6 11.9 11.3 11.3 11.4 12.6 11.0 9.5 10.7
Orbit diameter 4.9 6.6 6.3 5.8 6.0 6.2 5.4 5.8 5.5 5.2 4.9 5.3 5.3
Interorbital width 6.2 5.8 6.1 6.1 6.4 6.2 6.0 6.1 6.0 6.1 5.8 5.4 5.7
Caudal-peduncle
depth
11.2 10.4 11.2 10.5 11.4 10.9 11.3 11.5 11.6 10.9 11.1 10.9 11.1
Caudal-peduncle
length
9.8 11.1 10.3 9.6 9.3 7.9 9.5 9.8 10.3 9.8 9.4 10.3 9.4
Predorsal length 30.2 31.1 30.8 30.6 31.0 30.3 29.7 30.9 30.4 31.1 28.6 28.0 29.0
Preanal length 50.6 54.3 51.6 52.9 52.2 51.6 49.0 51.0 51.4 52.8 51.1 49.9 51.0
Prepelvic length 32.4 34.1 32.7 32.2 34.5 32.8 30.6 34.6 33.3 32.4 31.6 30.9 32.8
Base of dorsal fin 58.2 55.0 54.0 56.6 56.6 58.5 57.7 57.2 57.7 55.9 57.3 59.6 59.0
First dorsal-fin
spine
5.3 3.8 4.9 4.0 broken 5.6 4.2 5.3 5.6 4.3 4.6 4.3 3.6
Second dorsal-fin
spine
6.7 5.5 6.5 5.6 7.0 7.4 5.7 6.5 7.5 5.5 5.5 5.4 5.3
Last dorsal-fin
spine
10.6 9.3 10.4 9.5 9.9 10.9 9.1 9.8 10.7 9.8 8.6 8.6 9.3
Longest dorsal-fin
ray (number)
13.3 (10) 12.8 (10) 13.6 (10) 13.1 (10) 13.9 (10) 13.0 (11) 13.3 (9¼10) 12.3 (10) 13.7 (3) 13.4 (10) 12.1 (10) 12.0 (9) 13.5 (10)
Base of anal fin 40.8 35.1 38.2 38.0 39.8 40.6 41.3 39.6 41.1 40.2 40.3 40.1 40.3
First anal-fin spine 3.7 2.7 4.0 3.0 3.4 3.8 3.1 3.0 3.5 3.8 4.0 2.5 3.4
Second anal-fin
spine
5.7 5.5 5.9 5.4 4.9 broken 5.3 4.9 5.4 5.1 5.2 4.4 5.7
Third anal-fin spine 7.3 7.6 8.4 7.6 7.2 broken broken 7.3 8.1 6.8 7.1 5.8 6.9
Longest anal-fin
ray (number)
13.6 (4) 12.5 (4) 13.1 (2) 12.0 (3) 12.5 (4) 12.4 (5) 11.3 (4) 12.5 (3) 14.6 (5) 12.7 (4) 11.6 (3) 11.0 (3) 11.0 (3)
Caudal-fin length 23.4 22.7 24.3 23.2 21.7 21.9 22.6 22.7 22.0 22.5 21.0 20.4 21.0
Pectoral-fin length 13.9 14.8 15.5 15.2 16.3 14.8 14.1 14.8 15.6 13.7 14.0 14.8 14.5
Pelvic-spine length 8.9 7.9 9.4 8.0 8.2 9.3 8.2 8.1 8.8 8.1 8.1 7.7 7.9
Pelvic-fin length 13.6 13.1 13.9 13.0 13.8 14.1 12.9 12.8 13.3 14.1 12.9 12.3 12.4
Tea et al.—Revision of the Pseudojuloides elongatus complex 561
markings on head and nape become light tan. Females
uniformly tan; fins hyaline.
Habitat and distribution.—Pseudojuloides crux is known from
Western Australia, from the Houtman Abrolhos islands,
north to the Dampier Archipelago (Fig. 5). It inhabits rubble
zones with prominent kelp and other macroalgae growth at
depths ranging from 3–25 m.
Etymology.—The specific epithet refers to the most famous
constellation in the southern celestial hemisphere, the Crux
Constellation or Southern Cross. It acknowledges the species’
southern distribution, as well as the dark upper body with
numerous spots present in males, which is reminiscent of a
starry night. To be treated as a noun in apposition.
Pseudojuloides paradiseus, new species
urn:lsid:zoobank.org:act:B4053466-D069-491B-B9F8-
1652945AADCC
Paradise Pencil Wrasse
Standard Japanese name: Otohime-bera
Figures 2C, 8–10; Table 4
Leptojulis sp.: Masuda et al., 1975: 304 (Izu Oceanic Park,
Japan).
Pseudojuloides elongatus (non Ayling and Russell, 1977);
Randall and Randall, 1981: 56 (revision, material examined
from Japan); Parenti and Randall, 2000: 36 (checklist);
Nishiyama and Motomura, 2012: 122 (color photos; Izu
Oceanic Park, Japan); Kato, 2016: 95 (color photos; Izu
Oceanic Park, Japan)
Pseudojuloides sp. 4: Kuiter, 2010: 339 (color photos A–E; Izu
Oceanic Park, Japan).
Holotype.—(Figs. 8A–B) KPM-NI 43448, 103.0 mm SL male,
Izu Oceanic Park, Jogasaki-Kaigan, east coast of Izu Peninsula,
west of Sagami Bay, 15 m, algal reef, W. Takase, 8 May 2017.
Paratypes.—(Fig. 8C) KPM-NI 43449, 62.5 mm SL female,
collected with holotype; KPM-NI 37140, 74.6 mm SL male,
KPM-NI 32599, 43.2 mm SL female, KPM-NI 37035, 69.7 mm
SL male, KPM-NI 37137, 130.0 mm SL male, KPM-NI 33519,
111.7 mm SL male, KPM-NI 37138, 104.0 mm male, KPM-NI
33520, 98.1 mm SL male, KPM-NI 37139, 86.3 mm SL
female, all from same locality as holotype, except collected
on algal reef in 30 m, H. Masuda, 1 August 1990.
Diagnosis.—Pseudojuloides paradiseus belongs to the Pseudoju-
loides elongatus complex, species of which differ primarily
from all other species of Pseudojuloides in lacking median
predorsal scales and in having a large canine tooth on corner
of mouth. It shares overlapping or nearly overlapping
meristic counts and morphometric proportions with Pseudo-
juloides elongatus and Pseudojuloides crux, but differs in having
males that are bright yellowish orange to orange-pink, upper
body strongly overlain in black, with 3–5 rows of metallic
blue spots, and unmarked females that are reddish orange to
brick red.
Description.—For data and description that do not differ
between species of this complex, see description of Pseudoju-
loides elongatus. Upper procurrent caudal-fin rays 6 (two
paratypes with 5); lower procurrent caudal-fin rays 5 (one
paratype with 6); total caudal-fin rays 25 (two paratypes with
24, one paratype with 26); lateral line with 27 tubed scales,
posteriormost overlying hypural joint, with an additional large
tubed scale on caudal fin (one paratype with 26 tubed scales on
one side; one paratype with two tubed scales on caudal fin on
one side); scales below lateral line to origin of anal fin 9 (one
paratype with 8 scales on one side); gill rakers 15 (15–17).
Vertebrae 9þ16 (one paratype with only 15 apparent caudal
vertebrae, owing to fusion of preural 2 and preural 3 centra);
first vertebral centrum about half width of subsequent
centra, bearing a relatively short neural spine; single supra-
neural, inserted in first interneural space; first dorsal
pterygiophore bearing one supernumerary spine and one
serial spine, inserted along with second pterygiophore in
second interneural space (one paratype with first pterygio-
phore in first interneural space and next two pterygiophores
in second interneural space); remaining dorsal pterygio-
phores insert one per interneural space, except for 18
th
interneural space with two pterygiophores (19
th
interneural
space with two pterygiophores in one paratype; 18
th
with
three pterygiophores in one paratype); first two haemal
spines with enlarged secondary haemal arches (see Russell,
1988: fig. 23), first recurved towards second haemal spine;
first anal pterygiophore bears two supernumerary spines and
one serial spine, inserted anterior to first haemal spine;
subsequent anal pterygiophores each bears a serially associ-
ated segmented ray and inserted one per interhaemal space,
except for two pterygiophores in 11
th
interhaemal space
(four paratypes with two pterygiophores in 10
th
interhaemal
space); ribs present on vertebrae 3 through 9; epineurals
present on vertebrae 1 through 12 (11–13; Fig. 2C).
Body elongate, greatest depth 4.4 (4.4–5.3) in SL, width 2.1
(1.8–2.3) in greatest depth; head pointed, length 3.6 (3.2–3.4)
in SL; dorsal profile of head slightly convex; snout long, 2.8
(2.7–3.2) in HL; caudal peduncle short and narrow, least depth
2.5 (2.6–3.2) in HL, its length 2.9 (2.6–3.3) in HL. Mouth
small, terminal, upper jaw nearly reaching a point vertically
below anterior nostril. A pair of large, moderately projecting,
and slightly recurved canines present anteriorly in each jaw,
upper pair slightly flared and widened, lower pair curving
forward and fitting between upper canines when mouth
closed; a short row of 4–7 (upper jaw) and 5–7 (lower jaw)
irregularly placed chisel-like incisiform teeth along each side
of jaws; a canine tooth sometimes present posteriorly on
upper jaw of large specimens.
Scales cycloid, moderately large except those on thorax
and nape, which are much smaller; head naked; fins unscaled
except basal portion of caudal fin and base of pelvic fins.
Dorsal- and anal-fin spines progressively longer, last dorsal-
fin spine 2.8 (3.1–3.6) in HL and 1.2 (1.3–1.6) in longest soft
dorsal-fin ray, third anal-fin spine 3.5 (3.5–4.5) in HL and 1.5
(1.3–1.7) in longest soft anal-fin ray; caudal fin truncate to
slightly rounded in females and males; caudal fin relatively
small, 1.3 (1.2–1.4) in HL; pectoral fins small, 2.0 (2.0–2.2) in
HL; origin of pelvic fins below lower base of pectorals, their
length 2.0 (2.0–2.3) in HL. Morphometric values are
summarized in Table 4.
Coloration of males in life.—Based on color photographs of
specimens when freshly dead, and live individuals in the
field (Figs. 8A, 9, 10A–B): upper head and body bright orange
to pinkish orange; lower part of head yellow; head with a
network of metallic blue to lilac markings; interorbital and
upper part of snout with 3–4 lilac stripes, often sinuous and
562 Copeia 108, No. 3, 2020
reaching upper lip; a pair of stripes of the same color present
from behind upper orbit to upper edge of operculum; a
second pair of lilac stripes run obliquely from lower jaw and
lower orbit to lower part of operculum; upper part of nape
and sometimes narrow area before dorsal fin with additional
lilac stripe; iris bright yellow to orange, with blue submar-
ginal ring around pupil; pectoral-fin base with conspicuous
yellow to black band, edged distally in metallic blue; lower
part of abdomen bright orange to yellow; dorsal half of body
bright orange to pinkish orange, overlain with black, and
with 3–5 rows of metallic pink to spots; spots increasingly
broken ventrally; ventral half of body bright orange to yellow
Fig. 8. Specimens of Pseudojuloides paradiseus, new species, collected from Izu Oceanic Park, Jogasaki-Kaigan, east coast of Izu Peninsula, west of
Sagami Bay, Japan. (A) Freshly euthanized male holotype, KPM-NI 43448, 103.0 mm SL; (B) preserved male holotype, KPM-NI 43448, 103.0 mm SL;
(C) freshly euthanized female paratype, KPM-NI 43449, 62.5 mm SL. Photographs by H. Senou (A, C) and Y. K. Tea (B).
Tea et al.—Revision of the Pseudojuloides elongatus complex 563
without any obvious markings; dorsal fin bright orange with
anterior interspinous membrane spaces bright green, break-
ing up into oblique bands toward posterior three-quarters of
fin; distal margin of fin narrowly metallic blue; anal fin
similar to dorsal fin, but with irregular, oblique lilac to light
blue bands; distal margin of fin metallic blue; caudal fin
translucent yellow, sometimes with metallic blue spots
present near base; pelvic fins hyaline blue-gray, spines and
rays metallic blue, with two bright red spots present between
each intermembrane space; pectoral fins pinkish hyaline.
Coloration of females in life.—Based on color photographs of
specimens when freshly dead, and live individuals in the
field (Figs. 8C, 10C): head and upper body uniformly reddish
orange to brick red, gradually fading to yellow-orange
ventrally; interorbital, nape, and predorsal area sometimes
dusky brown; iris bright yellow to orange, with blue
submarginal ring around pupil; dorsal, anal, and caudal fin
hyaline yellow; pelvic and pectoral fins translucent gray.
Coloration in preservative.—(Fig. 8B) Head and body tan;
black markings remain; pectoral fin axil spot becomes black;
metallic blue spots on back become pale tan; nape dusky;
sinuous markings on head and nape become light tan.
Females uniformly tan; fins hyaline.
Distribution and habitat.—Pseudojuloides paradiseus is sporad-
ically distributed in the following localities of southern Japan
(Fig. 5): Izu-Oshima (KPM-NR 94350) and Miyake Island, the
Izu Islands; Tateyama Bay, east of Sagami Bay; the Izu
Oceanic Park, east coast of the Izu Peninsula, Sagami Bay;
Osezaki, west of the Izu Peninsula (KPM-NR 22025), Suruga
Bay; Kushimoto, Wakayama Prefecture, southern end of the
Kii Peninsula; Kashiwa Island, Shikoku; Mishima Island,
Fig. 9. Males of Pseudojuloides paradiseus, new species: (A) male from Okino-shima Island, Munakata, Fukuoka, northwestern Kyushu; (B) male,
approximately 150 mm TL, underwater photo in 15 m, Izu Oceanic Park, Shiuoka Prefecture, Japan. Photographs by Y. Yogo and K. Nishiyama,
respectively.
564 Copeia 108, No. 3, 2020
Yamaguchi Prefecture; Myoken-ura, Amakusa, Kuwamoto
Prefecture; Yunomoto, Iki Island, Nagasaki; Okino-shima
Island, Munakata, Fukuoka; and Yakataishi, Karatsu, Saga
Prefecture (Yogo, 2000; Shimada, 2013; present study). It
appears to inhabit rubble zones with prominent kelp and
macroalgae growth, at depths ranging from 4–40 m.
Etymology.—The specific epithet is the Latin for paradise, in
reference to the arresting live coloration of the males.
Comparisons and discussion of the Pseudojuloides elon-
gatus complex
Species of the genus Pseudojuloides can be distinguished
from all other labrids by the following combination of
characters: body slender and elongate; dorsal-fin rays IX,11
or XI,12; anal-fin rays 12 or 13; lateral line continuous, with
26–28 pored scales, the last overlapping the hypural crease;
mouth small, the side of jaws with chisel-like incisiform
Fig. 10. Underwater photographs of Pseudojuloides paradiseus, new species, from Izu Oceanic Park, Shizuoka Prefecture, Japan: (A) male,
approximately 150 mm TL, photographed in 15 m; (B) male, approximately 150 mm TL, photographed in 15 m; (C) female, approximately 100 mm
TL, photographed in 12 m. Photographs by K. Nishiyama.
Tea et al.—Revision of the Pseudojuloides elongatus complex 565
teeth. Randall and Randall (1981) distinguished Pseudoju-
loides elongatus from the rest of the genus based on the
following combination of characters: modal dorsal-fin rays
IX,12; modal pectoral-fin rays 12; no median predorsal scales;
5–7 suborbital pores; corner of mouth with a canine tooth;
and membranes of spinous portion of dorsal fin less than
10% higher than spine tips. Our examined specimens of
Pseudojuloides crux, P. elongatus, and P. paradiseus agree well
with this diagnosis. Previous molecular phylogenetic studies
of the genus based on mitochondrial COI suggest that P.
elongatus is fairly divergent from the other species of
Pseudojuloides, occupying a separate lineage from the other
more tropical species (Victor and Edward, 2016). Our results
indicate that Pseudojuloides elongatus,P. crux, and P. paradiseus
are likely to form a monophyletic group, hereafter referred to
as the Pseudojuloides elongatus complex.
While the three species are readily separated on the basis of
coloration differences (summarized in Table 5), their rela-
tionships inferred on the basis of molecular sequence data are
more problematic, largely exacerbated by the lack of
comparative genetic material. All pairwise divergences were
less than 1% at COI and 16S. While this intraspecific level of
divergence is low, it is not uncommon in coral reef fishes,
particularly for many species of sexually dimorphic labrids,
where low levels of genetic divergence are observed despite
large differences in coloration and other morphological
phenotypes (Victor and Randall, 2014; Allen et al., 2016;
Tea et al., 2019b). These traits are hypothesized to have
evolved more rapidly at specific loci under sexual selection,
compared with other regions of the genome.
The sharing of control region haplotypes between the
individuals of P. crux and P. elongatus is equally problematic.
Table 4. Morphometric values for Pseudojuloides paradiseus expressed as percentage SL.
Holotype Paratypes
KPM-NI
43448
KPM-NI
32599
KPM-NI
43449
KPM-NI
37035
KPM-NI
37140
KPM-NI
37139
KPM-NI
33520
KPM-NI
37138
KPM-NI
33519
KPM-NI
37137
Standard length (mm) 103.0 43.2 62.5 69.7 74.6 86.3 98.1 104.0 111.7 130.0
Greatest body depth 22.8 19.0 20.3 18.8 21.0 20.2 20.5 22.1 21.7 22.7
Body depth at dorsal
origin
20.7 18.8 17.9 17.1 19.0 18.3 19.8 19.9 20.6 21.6
Body width 10.9 8.8 9.6 10.2 9.5 10.1 10.6 12.0 9.2 11.1
Head length 28.6 31.5 30.1 29.3 30.4 30.5 30.4 29.7 29.5 30.3
Snout length 10.2 10.0 10.9 10.0 10.5 11.2 10.7 10.9 10.7 10.9
Orbit diameter 5.0 6.9 6.7 5.5 6.1 5.4 5.0 4.9 4.7 4.4
Interorbital width 5.6 5.8 6.1 5.6 5.6 6.1 6.0 5.9 6.1 6.2
Caudal-peduncle depth 11.5 9.7 11.5 10.2 11.8 10.8 10.9 11.2 11.5 11.4
Caudal-peduncle length 10.0 10.9 9.8 11.0 9.2 9.2 9.8 9.2 9.3 10.5
Predorsal length 27.3 30.6 28.0 27.1 29.4 29.3 28.6 28.9 28.2 29.1
Preanal length 50.6 51.6 52.5 49.8 49.3 53.1 50.2 50.3 52.3 49.8
Prepelvic length 32.6 31.7 32.6 31.1 32.2 32.0 31.5 31.3 31.8 30.3
Base of dorsal fin 59.6 55.6 56.8 57.8 59.2 57.1 62.1 58.9 58.0 58.2
First dorsal-fin spine 5.1 3.9 3.7 4.0 4.2 4.3 3.6 4.1 4.8 4.7
Second dorsal-fin spine 6.9 5.6 4.5 broken 5.5 5.8 4.9 6.3 6.4 6.6
Last dorsal-fin spine 10.9 9.3 8.5 9.5 8.7 8.7 9.3 8.3 9.6 9.6
Longest dorsal-fin ray
(number)
12.1 (9) 13.9 (9) 11.8 (9) 13.6 (9) 13.7 (9) 12.7 (9) 13.6 (9) 12.6 (9) 12.7 (9) 13.4 (9)
Base of anal fin 41.7 38.7 38.6 39.2 39.7 37.9 38.5 39.8 41.0 39.7
First anal-fin spine 3.4 3.5 2.7 3.0 2.9 2.9 2.8 3.0 3.0 2.9
Second anal-fin spine 5.6 6.9 5.3 5.9 5.8 5.3 4.8 5.2 5.1 4.6
Third anal-fin spine 8.3 9.0 7.8 8.2 8.4 7.5 7.1 6.8 7.3 broken
Longest anal-fin ray
(number)
12.2 (7) 11.8 (2) 11.5 (2) 11.9 (3) 12.1 (3) 11.8 (3) 11.9 (3) 11.6 (4) 12.3 (3) broken
Caudal-fin length 22.0 25.5 23.8 23.4 23.5 21.1 21.7 21.9 21.8 broken
Pectoral-fin length 14.7 15.0 14.1 14.2 15.0 14.6 14.0 13.9 broken 14.2
Pelvic-spine length 9.3 8.6 8.8 8.3 8.7 8.3 8.3 8.3 8.6 7.9
Pelvic-fin length 14.0 13.4 13.1 13.5 13.9 13.1 12.9 12.8 12.8 13.2
Table 5. Summary of live coloration characters for selected species of Pseudojuloides.
Pseudojuloides elongatus Pseudojuloides crux Pseudojuloides paradiseus
Ground coloration of males Pinkish orange to olivaceous Greenish yellow to yellowish orange Yellowish orange to orange-pink
Markings on upper body
of males
Upper body without black,
sometimes with 0–4 rows
of metallic blue spots
Upper body overlaid with black, with
3–5 rows of metallic blue stripes
breaking into spots
Upper body overlaid with black, with
3–5 rows of metallic blue stripes
breaking into spots
Ground coloration of females Pinkish orange to olivaceous Olivaceous Reddish orange to brick red
566 Copeia 108, No. 3, 2020
This phenomenon is frequently observed in lineages that
have either not completely sorted, and so ancestral poly-
morphisms are still retained in some individuals (Degnan
and Rosenberg, 2009), or in introgressed lineages that are
hybridizing. Differentiating incomplete lineage sorting from
introgression is difficult without larger amounts of genomic
data, although introgression is unlikely to occur in disjunct,
allopatric populations such as these where contemporary
gene flow is not likely to occur. Indeed, comparisons of
individuals of P. elongatus vs. P. paradiseus, and of P. crux vs. P.
paradiseus, show a difference of 2.1% in the mitochondrial
control region, suggesting that the haplotype sharing
between the individuals of P. crux and P. elongatus is likely
to be a result of incomplete lineage sorting.
The multispecies-coalescent model implemented in ASTRAL
takes into account the possibility of incomplete lineage sorting
by reconstructing a species tree from a set of gene trees. Our
results show support for a more divergent Western Australian
lineage and a more closely related southern Japan þsouthwest-
ern Pacific lineage, consistent with our PCA of morphological
variables. These results indicate the presence of cryptic diversity
and early stages of diversification between populations.
Indeed, the three species exhibit strong, consistent differ-
ences in coloration patterns corresponding to their biogeo-
graphical distributions that are suggestive of reproductive
isolation. Moreover, unlike other species of Pseudojuloides
(with the exception of Pseudojuloides atavai), where females are
remarkably similar in coloration and are otherwise difficult to
separate in the field, the three species of the Pseudojuloides
elongatus complex have females that are easily differentiated
from each other with regard to overall coloration.
Perhaps the most unusual aspect of the Pseudojuloides
elongatus complex is in their ecology. Unlike other members
of the genus, nearly all of which are associated with deeper,
tropical coral reefs, members of the Pseudojuloides elongatus
complex are frequently observed in relatively shallow, sub-
tropical habitats dominated by kelp and macroalgae. One
additional taxon of interest here is the elusive western Indian
Ocean endemic Pseudojuloides argyreogaster (Fig. 11), which is
known only from few specimens collected from shallow
seagrass beds (Victor and Randall, 2014). Randall and Randall
(1981) commented on a potentially close relationship shared
between P. argyreogaster and P. elongatus (¼P. elongatus com-
plex), but noted that P. argyreogaster differed from P. elongatus
in having 3–5 more suborbital pores, a higher interspinous
membrane height, and one fewer lower procurrent caudal-fin
ray. Owing to the elusive nature of this species and the lack of
comparative genetic material, we are unable to confidently
assign Pseudojuloides argyreogaster to the Pseudojuloides elonga-
tus complex. However, given that the characters separating it
from Pseudojuloides elongatus are prone to variation, it is likely
that the four species form a monophyletic group, but further
study is required to investigate this relationship.
Anti-equatorial patterns in coral reef taxa
The biogeographical distributions for the three species in the
Pseudojuloides elongatus complex conform to an anti-equatorial
biogeographic pattern as defined by Randall (1981), where taxa
are distributed in both hemispheres but are absent from
equatorial latitudes (Fig. 5). The geographical components of
the Pseudojuloides elongatus complex consist of Western Austra-
lian, eastern Australian/southwest Pacific, and East Asian areas.
The same general patterns have been reported for various other
marine taxa, including a labrid clade consisting of Pseudolabrus
and Notolabrus (Mabuchi et al., 2004), several species from the
genus Macropharyngodon (M. kuiteri þM. moyeri þM. vivienae;
pers. obs.), and the microcanthid genus Microcanthus (Tea et al.,
2019a; Tea and Gill, 2020). Of these taxa, Microcanthus shares a
nearly identical distribution pattern to the P. e l o n g a t u s complex,
but with an additional population in Hawaii.
Comparison of area relationships and population-level
divergences between Microcanthus and the Pseudojuloides
elongatus complex, however, reveals some noteworthy differ-
ences. Unlike the latter, where divergence at the population
level is reflected largely by differences in coloration, Micro-
canthus exhibits deep genetic divergences at the population
level with little differences in coloration (Tea et al., 2019a).
Molecular dating and historical biogeographical analyses of
Microcanthus suggest a dispersal event coinciding with the
last Pleistocene glacial cycles (Tea et al., 2019a), allowing
enough time for genetic separation between isolated popu-
lations. However, no such levels of genetic divergences were
observed between the various species of the Pseudojuloides
elongatus complex (although more extensive sampling is
needed to assess this more confidently).
The contrasting patterns between area relationships and
levels of genetic differentiation in the Pseudojuloides elongatus
complex versus Microcanthus suggest that biogeographical
patterns among anti-equatorial taxa may be more complex
than previously assumed. Similarities in contemporary distri-
bution patterns may not necessarily reflect similarities in
Fig. 11. Underwater photograph of
Pseudojuloides argyreogaster from
Seychelles. Note the green body
coloration and blue facial markings
similar to those in Pseudojuloides
elongatus. Photograph by R. Daly.
Tea et al.—Revision of the Pseudojuloides elongatus complex 567
historical processes, as evidenced by differences in rates of
mutation between taxa, selective drivers for speciation, and
time-dependent dispersal and biogeographical processes. Until
a rigorous study can be performed on the Pseudojuloides
elongatus complex, we are unable to comment beyond alluding
to similarities in anti-equatorial patterns between these taxa. A
more comprehensive study should consider the potentially
related Pseudojuloides argyreogaster, inclusion of which would
make the Western Indian Ocean an additional area component
in the biogeography of this species complex. In at least one
other labrid group (Macropharyngodon), this region appears to be
of importance in considering historical processes leading to
contemporary anti-equatorial distributions. Both sexes of
Macropharyngodon kuiteri, M. moyeri,andM. vivienae share
coloration patterns that depart from those of congeners, and
the three species likely form a monophyletic group. However,
only one of the species (M. kuiteri) was included in the
phylogenetic study of Macropharyngodon by Read et al. (2006),
thus the proposed relationship for these species should be
regarded as tentative. The Pacific species M. kuiteri and M.
moyeri have an anti-equatorial distribution in southern Japan
and the southwest Pacific, respectively, while M. vivienae is
sporadically distributed along the southeastern coast of Africa
and the Mascarenes. Whether this Western Indian Ocean þ
anti-equatorial Japan and southwest Pacific Ocean pattern is
reflected in the relationships of these species (and by extension
Pseudojuloides), requires further investigation.
DATA ACCESSIBILITY
The mitochondrial and nuclear S7 sequence alignment used
to infer the gene trees and species tree in this study and the
supplemental appendices are available at https://www.
copeiajournal.org/ci-19-316. GenBank and BOLD accession
numbers are included in the alignment and Supplemental
Appendix B (MK120502–12; MN076190–93).
ACKNOWLEDGMENTS
We thank M. McGrouther, A. Hay, S. E. Reader, G. Moore, and
M. Allen for curatorial assistance and the loan of specimens.
The holotype and one paratype of Pseudojuloides paradiseus
were collected by W. Takase. We thank J. Gill who collected
and graciously donated one specimen of Pseudojuloides crux for
this study (paratype; AMS. I.48996-001). We thank I. Shaw, J.
Sear, R. H. Kuiter, G. R. Allen, J. Gill, R. Daly, and K. Nishiyama
for providing excellent photographs used in this study and A.
Hay and S. E. Reader for X-radiographs. We thank B. C. Victor
and A. Teitelbaum who provided tissue samples of Pseudoju-
loides elongatus from New Caledonia. We thank B. Frable, W. B.
Ludt, and S. Ho who provided helpful comments that greatly
improved a previous version of this manuscript.
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Appendix 1. Percentage difference in uncorrected pairwise distances for mitochondrial COI, 16S, and control region sequences for selected
individuals of Pseudojuloides elongatus (sensu lato).
COI 16S Control region
Southwest
Pacific Ocean
Southern
Japan
Western
Australia
Southwest
Pacific Ocean
Southern
Japan
Western
Australia
Southwest
Pacific Ocean
Southern
Japan
Western
Australia
Southwest Pacific Ocean — — —
Southern Japan 0.5 — 0.2 — 2.1 —
Western Australia 0.9 0.4 — 0.4 0.2 — 0 2.1 —
Appendix 2. Percentage difference in uncorrected pairwise distances for nuclear S7 and RAG2 sequences for selected individuals of Pseudojuloides
elongatus (sensu lato). N/A indicates missing data.
S7 RAG2
Southwest
Pacific Ocean
Southern
Japan
Western
Australia
Southwest
Pacific Ocean
Southern
Japan
Western
Australia
Southwest Pacific Ocean — —
Southern Japan 0.6 — N/A —
Western Australia 1.3 1.2 — N/A 1.3 —
Tea et al.—Revision of the Pseudojuloides elongatus complex 569
