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Two New Species of Pencil Wrasses (Teleostei: Labridae: Pseudojuloides)from
Micronesia and the Marquesan Islands
Yi-Kai Tea
1,2,¶
, Brian D. Greene
3,¶
, John L. Earle
3
, and Anthony C. Gill
1,2,4
Pseudojuloides pluto, new species, is described on the basis of the holotype and 11 paratypes from Wake Island,
northeastern Micronesia, and nine paratypes from the Maug Islands, Northern Mariana Islands. The new species has
previously been confused with Pseudojuloides atavai, but molecular analysis of mitochondrial COI reveals a difference of
6.8% in sequence data between both species, in addition to differences in meristic, morphometric, and coloration
details. A second new species, Pseudojuloides proserpina, is described from Fatu Hiva, Marquesan Islands on the basis of
the male holotype. The two new species are closely allied to Pseudojuloides atavai, and together form a species complex
that differs from other members of the genus in having males that share the following combination of characters:
interspinous membrane between the anterior two to three spines of the dorsal fin with a black spot; head extensively
reticulate (reduced in P. pluto, new species); dorsal-fin base with a pink stripe; abdominal region behind pectoral and
pelvic fins pale lilac to orangey pink (width of this region dependent on species) with a crosshatch or honeycomb
pattern; and extensive black coloration over at least posterior half of body. Additionally, females of both P. atavai and P.
pluto, new species, are distinctly bicolored (versus unicolored and suffused in all other congeneric species). Although
the female form of P. proserpina, new species, is not known, it is likely that it shares this general coloration pattern,
which may serve as an additional character uniting members of the Pseudojuloides atavai complex. We briefly discuss the
phylogenetic relationships of Pseudojuloides inferred on the basis of mitochondrial DNA.
THE labrid fish genus Pseudojuloides includes mostly
tropical to subtropical species found throughout the
Indo-Pacific. Randall and Randall (1981) revised the
genus, recognizing eight species, five of which were new.
Since then, a further eight new species have been described,
bringing the number of valid species within the genus to 18
(Tea et al., 2020, in this volume). With the exception of
Pseudojuloides argyreogaster and the Pseudojuloides elongatus
complex (Tea et al., 2020, in this volume), species of
Pseudojuloides are typically associated with coral reefs or
rubble zones adjacent to coral cover. The genus is unique
among labrids in having chisel-like incisiform teeth and
slender fusiform bodies, lending to common names of
slender or pencil wrasses. Like many other labrids, terminal
phase males are typically more brightly colored than the
females, which for species of Pseudojuloides are usually
unmarked and uniformly colored or suffused with color.
Of these, Pseudojuloides atavai is unusual among congeners
in having attractively patterned females that are distinctly
bicolored. The species is primarily known from French
Polynesia and surrounding islands in southeastern Oceania.
However, reports of Pseudojuloides atavai occurring outside of
this region have been documented in the literature by several
authors, particularly in the Mariana and Ogasawara Islands
(Springer, 1982; Myers, 1988; Myers and Donaldson, 2003;
Kato, 2016). This is due partly to the distinctive coloration
pattern of the females, which no other species of Pseudoju-
loides resemble. On the basis of differences in male coloration
patterns, Kuiter (2010) recognized those from the Mariana
Islands as a putative undescribed species; however, no
systematic work has been done to address the taxonomic
status of these population, and whether or not Pseudojuloides
atavai is truly a single, widespread species.
Morphological and molecular comparisons of specimens
identified as Pseudojuloides atavai from northeastern Micro-
nesia indicate that they are different from the nominal P.
atavai from French Polynesia. Accordingly, we describe
Pseudojuloides pluto, new species, on the basis of 21 specimens
collected from Wake Island, northwestern Micronesia, and
the Maug Islands, Northern Mariana Islands. Additionally,
we describe a second species, Pseudojuloides proserpina, new
species, on the basis of the male holotype collected from Fatu
Hiva, Marquesan Islands. Previous phylogenetic estimates
inferred on the basis of mitochondrial DNA have shown
support for a highly divergent Pseudojuloides atavai with
respect to other members of the genus (Victor, 2017). We
provide a brief account of P. atavai with new observations
based on examination of two paratypes and discuss the
phylogenetic relationships of Pseudojuloides inferred on the
basis of mitochondrial DNA.
MATERIALS AND METHODS
Meristics, morphometrics, and specimen deposition.—Methods
for counting and measuring follow Randall and Randall
(1981). Measurements were recorded to the nearest 0.1 mm
using digital calipers. All measurements to the tip of the
snout were made to the midanterior part of the upper lip.
Length of specimens are given in mm standard length (SL),
which was measured from the snout tip to the middle of the
caudal peduncle at the vertical through the posterior edge of
the dorsal hypural plate. Head length (HL) was measured
from the snout tip to the posteriormost edge of the opercular
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, 1 William Street, Sydney, New South Wales 2010, Australia.
3
Association for Marine Exploration, 420 N Kalaheo Avenue, Kailua, Hawaii 96740.
4
Macleay Collections, Chau Chak Wing Museum, The University of Sydney, New South Wales 2006, Australia.
¶
Authors contributed equally.
Submitted: 8 November 2019. Accepted: 11 April 2020. Associate Editor: M. T. Craig.
Ó2020 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-19-327 Published online: 15 October 2020
Copeia 108, No. 3, 2020, 679–691
membrane. Snout length was measured over the shortest
distance from the snout tip to the orbital rim. Orbit diameter
was measured as the horizontal width between the fleshy
edges of the orbital rim. Interorbital width is the least bony
width. Predorsal, preanal, and prepelvic lengths were
measured from the snout tip to the base of the first spine
of the relevant fin. Body width was measured at the bases of
the pectoral fins. Caudal peduncle length was measured from
the base of the last anal-fin ray to the ventral edge of the
caudal fin at the vertical though the posterior edge of the
ventral hypural plate. Caudal peduncle depth was measured
where least. Caudal-fin length was measured horizontally
from the hypural crease to the tip of the longest ray.
Measurements of fin spines were from the proximal base of
the spine to the distal tip, excluding any filamentous
extensions. Pectoral-fin length was measured as the length
of the longest ray. Pelvic-fin length was measured from the
base of the spine to the distal tip of the longest ray. Total
number of lateral-line scales include the last enlarged pored
scale on the caudal fin, posterior to hypural crease.
Morphometric values are summarized in Table 1, expressed
as percentage SL.
The last ray in the dorsal and anal fins is divided at its base
and was counted as a single ray. 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 (pu2). Counts of principal and
branched caudal-fin rays are presented in the form upper þ
lower, 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 lateral line to base of first dorsal-fin spine 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.
Table 1. Morphometric values for Pseudojuloides pluto, new species, and P. proserpina, new species, expressed as percent SL. An expanded version
of this table with values for each paratype is available as supplemental material (see Data Accessibility).
Pseudojuloides pluto, new species
Pseudojuloides
proserpina,
new species
HolotypeHolotype
Paratypes
Number Range
Mean 6Standard
Deviation jNumber
Sex Male 12 Female j8 Male Male
Standard length (in mm) 78 20 54.5–93.5 69.9612.5 91
Greatest body depth 26.8 20 22.4–27.1 24.161.3 28
Body depth at dorsal-fin origin 24.7 20 20.8–24.8 22.760.9 24.9
Body width 11.9 20 10.1–13.0 11.660.8 11.6
Head length 31.9 20 28.7–33.1 30.461.1 29.7
Snout length 10.3 20 9.0–11.9 10.360.7 9.8
Orbit diameter 6 20 5.1–7.7 6.460.7 5.5
Interorbital width 6.4 20 6.0–7.4 6.760.4 6.5
Caudal-peduncle depth 11.4 20 10.1–11.8 11.260.5 12
Caudal-peduncle length 11.8 20 9.0–13.1 10.561.1 9.9
Predorsal length 31.5 20 29.6–33.6 31.661.3 30
Preanal length 52.1 20 31.0–55.5 50.568.3 56.7
Prepelvic length 33.2 20 31.8–36.0 34.361.0 33.8
Base of dorsal fin 57.7 20 55.6–59.3 57.661.1 59.7
First dorsal-fin spine length 6.7 20 5.4–7.4 6.160.6 5.9
Second dorsal-fin spine length 8.6 20 6.6–8.9 7.760.7 6.5
Last dorsal-fin spine length 10.3 20 9.0–10.8 9.960.5 8.8
Longest dorsal-fin ray length 12.1 20 10.2–13.2 11.460.8 10.3
Longest dorsal-fin ray (ray number) 9
th
20 9
th
–10
th
9
th
1
st
Base of anal-fin length 37.4 20 33.8–38.9 36.161.5 35.4
First anal-fin spine length 3.1 20 2.2–3.9 2.860.4 3
Second anal-fin spine length 4.7 20 4.1–6.3 5.060.6 6
Third anal-fin spine length 7.6 20 7.0–8.9 7.860.6 7.4
Longest anal-fin ray length 12.4 19 9.9–11.9 10.860.6 9.6
Longest anal-fin ray (ray number) 3
rd
19 3
rd
–4
th
3
rd
3
rd
Caudal-fin length 20.9 17 18.9–21.8 20.460.9 19
Pectoral-fin length 16.3 20 13.6–18.1 15.961.0 16.5
Pelvic-spine length 9.7 20 8.3–10.4 9.260.5 8.8
Pelvic-fin length 15.4 20 12.7–15.8 14.360.8 14
680 Copeia 108, No. 3, 2020
Terminology of intermuscular bones follows Patterson and
Johnson (1995) and Johnson and Patterson (2001).
In the descriptions of the new species, data are presented
for the holotype, followed by data for the paratypes in
parentheses, where variation was noted. 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. Type specimens are deposited in the Bernice Pauahi
Bishop Museum, Honolulu (BPBM) and the Australian
Museum, Sydney (AMS). We made use of photographs from
the National Museum of Natural History, Smithsonian
Institution (USNM), Royal Ontario Museum (ROM), as well
as those from the Image Database of Fishes housed at the
Kanagawa Prefectural Museum of Natural History, Odawara
(KPM-NI). Photographs from the latter are assigned unique
numbers with the prefix KPM-NR. Note that owing to zero-
padding, a seven-digit number is used for the catalogue
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. Institutional codes
follow Sabaj (2019).
Taxon sampling, sequencing, and phylogenetic analysis.—This
study includes 17 ingroup species of Pseudojuloides and
Leptojulis cyanopleura as an outgroup. Comparative species
of Pseudojuloides were selected following Victor (2017).
Publicly available sequences were obtained from GenBank.
Tissue samples of the new species were obtained from the
right pelvic or pectoral fin, preserved in 100% ethanol, and
stored at –208C prior to extraction. DNA was extracted using
the DNeasy Blood and Tissue kit (Qiagen) following the
manufacturer’s protocol and sequenced de novo. Mitochon-
drial 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 Macrogen
(Seoul, South Korea). Forward and reverse contigs were
aligned and trimmed separately using GENEIOUS Prime
2019.1.1 (Biomatters). Sequences were then combined with
those available from GenBank and aligned using the
MUSCLE v3.8.31 algorithm (Edgar, 2004). The final sequence
alignment consisted of 652 base pairs. Accession numbers for
the new species and comparative sequences of other
Pseudojuloides are presented in the Supplemental Appendix
(see Data Accessibility).
We analyzed the molecular data set using both Bayesian
inference and maximum likelihood. The best fitting substi-
tution model was selected using PartitionFinder2 (Lanfear et
al., 2016). Bayesian analyses were conducted in MrBayes
v3.2.5 (Ronquist et al., 2012) using Markov chain Monte
Carlo sampling, with one cold and three heated Markov
chains. Samples were recorded every 5310
3
generations over
a total of 5310
7
generations, with the initial 25% of samples
discarded as burn-in. The analysis was run in duplicate and
checked for convergence in the program Tracer v1.7.1
(Rambaut et al., 2018). TreeAnnotator v2.2.0 (Bouckaert et
al., 2014) was used to generate the maximum-clade-credibil-
ity-tree. Maximum likelihood analyses were conducted in
RAxML v8.2.12 (Stamatakis, 2014). The analysis was per-
formed in duplicate to check for local optima, each using ten
random starts. Bootstrapping was performed using 1,000
pseudoreplicates of the data.
Pseudojuloides atavai
Polynesian Wrasse
Figures 1, 2A
Pseudojuloides atavai (Randall and Randall, 1981): 58, figs. 4, 5
(type locality: off West Harbor [sic; ¼Western Harbour],
Pitcairn Island); Springer, 1982: 129 (in part, distribution
pattern; southeast Oceania); Randall, 1999: 22 (checklist of
fishes collected from the Pitcairn Islands); Kuiter, 2010:
337 (color photograph A–C, E; live photographs of
specimens from Rarotonga, Cook Islands, South Pacific
Ocean, and aquaria).
Diagnosis.—The following combination of characters distin-
guishes Pseudojuloides atavai from congeners: body elongate
(greatest body depth 22.6–25.3% SL); dorsal-fin rays IX,11;
anal-fin rays III,12; pectoral-fin rays 13; tubed lateral-line
scales 28; interspinous membrane between anterior two to
three spines of dorsal fin with a black spot; head yellow and
extensively reticulate in males; abdominal region behind
pelvic and pectoral fins orange to orange-pink with yellow
crosshatch markings; posterior body purple black; females
distinctly bicolored, orange-brown dorsally, white ventrally.
Remarks.—A relatively large species (maximum size 115.8
mm SL; Randall and Randall, 1981), P. atavai is known from
oceanic reefs in the southeastern Pacific Ocean. In French
Polynesia, it is reported from the Society Islands (Fig. 1B, D,
E), Tuamotu Islands, Rapa, and the Austral Islands. Within
the northern Tuamotus, it has been reported from Takaroa
and Temoe (Randall and Randall, 1981); and in the Austral
Islands, from Rurutu (Fig. 1A), Tabuai (Fig. 1C), and ˆ
Iles Maria
(Fig. 1F). It also occurs in the Pitcairn Islands, where it has
been observed at Pitcairn (type locality), Ducie, and Oeno
(paratype; AMS I.21716-001). The species also occurs in
Raratonga, Cook Islands (Kuiter, 2010). It frequents outer
reefs away from lagoons and sheltered bays at depths
between 12–30 m. Records of this species occurring from
the Marianas and Ogasawara Islands in the Western Pacific
Ocean are of a different species (see below).
We add the following new observations: scales cycloid;
vertebrae 9þ16; first vertebral centrum about half width of
subsequent centra, bearing relatively short neural spine;
single supraneural, 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; remaining dorsal pterygiophores
insert one per interneural space, except for the 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;
subsequent anal pterygiophores each bearing a serially
associated segmented ray and insert 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; pu2 haemal spine
autogenous; 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).
Tea et al.—Two new species of Pseudojuloides 681
Etymology.—The epithet atavai is Tahitian for pretty, in
reference to the attractive coloration of both sexes of the
species (Randall and Randall, 1981).
Material examined.—AMS I.21716-001, paratype, 89.1 mm
SL, north side of Oeno Atoll, Pitcairn Islands, outer reef, coral
rubble and coral, 25 m, spear, J. E. Randall, 19 December
1970; AMS I.21717-001, paratype, 66.4 mm SL, off The Rope,
Pitcairn Island, outer reef, coral rubble and coral, 28–31 m,
rotenone, J. E. Randall, D. B. Cannoy, and S. R. Christian, 23
December 1970.
Pseudojuloides pluto, new species
urn:lsid:zoobank.org:act:3C3F4AE6-C7B1-4AE6-8B7D-EC-
BA1B50C6B0
Narcissus Pencil Wrasse
Figures 3–5, Table 1
Pseudojuloides atavai (non Randall and Randall, 1981):
Springer, 1982 (in part, distribution pattern; Guam);
Myers, 1988 (checklist; Mariana Islands); Lobel and Lobel,
2004: 74 (checklist; Wake Atoll, Micronesia); Kuiter, 2010:
337 (color photograph D; Saipan, Northern Mariana
Archipelago); Kato, 2016: 99 (color photographs; Ogasa-
wara Islands, Japan).
Pseudojuloides sp. A: Myers and Donaldson, 2003 (checklist;
Saipan).
Pseudojuloides sp. 2: Kuiter, 2010: 334 (color photographs A–
C; Saipan).
Holotype.—BPBM 40392, 78.0 mm SL, south side of Wake
Island, 11–20 m, patch reef on sand rubble bottom, hand
nets, J. L. Laughlin, 15 October 2005 (Fig. 3A, D).
Paratypes.—BPBM 41375, 6, 63.2–88.4 mm SL (collected with
holotype; Fig. 3F); BPBM 40391, 9, 54.5–88.9 mm SL,
Northern Mariana Islands, Maug Islands, 11 m, sand and
rubble bottom, hand nets, J. L. Laughlin, 13 September 2005
(Fig. 3B, C, E); AMS I.49040-001, 5, 57.8–93.5 mm SL, west
end of Wake Island, 19.28978N, 166.60988E, 15–20 m, patch
reef on sand rubble bottom, hand nets, B. D. Greene, 29 July
2019.
Diagnosis.—The following combination of characters distin-
guishes Pseudojuloides pluto from congeners: body elongate
(greatest body depth 22.4–27.1% SL); dorsal-fin rays IX,11;
anal-fin rays III,12; pectoral-fin rays 13–14 (usually 13); tubed
lateral-line scales 27–29 (usually 28); interspinous membrane
between anterior two to three spines of dorsal fin with an
irregular black spot; males olive green dorsally, fading to
whitish green ventrally, colors usually entirely obscured by
black coloration; nape and upper orbit with pinkish red stripe,
continuing along base of dorsal fin to upper edge of caudal
peduncle; abdominal region behind pectoral and pelvic fins
with a narrow lilac to orangey pink bar, about width of orbit,
from dorsal-fin origin to pelvic fin; females distinctly
bicolored, dorsal half orange-brown suffused with black.
Description.—Dorsal-fin rays IX,11, all segmented rays
branched (one paratype with first ray unbranched); anal-fin
Fig. 1. Freshly euthanized specimens of Pseudojuloides atavai: (A) male, USNM 424042, 121.6 mm SL, Rurutu, Austral Islands, French Polynesia;
(B) female, USNM 392362, 83.8 mm SL, Moorea, Society Islands, French Polynesia; (C) male, USNM 424098, 121.0 mm SL, Tubuai, Austral Islands,
French Polynesia; (D) female, USNM 392362, 83.8 mm SL, Moorea, Society Islands, French Polynesia; (E) male, ROM 59428, 86.4 mm SL, Moorea,
Society Islands, French Polynesia; (F) female, USNM 423273, 66.8 mm SL, Maria Atoll, Austral Islands, French Polynesia. Photographs by J. T. Williams
(A–D, F) and R. Winterbottom (E).
682 Copeia 108, No. 3, 2020
rays III,12, all segmented rays branched; pectoral-fin rays 13
(one paratype with 13/14), upper two unbranched; pelvic-fin
rays I,5; principal caudal-fin rays 7þ7; upper and lowermost
unbranched; upper procurrent caudal-fin rays 6; lower
procurrent caudal-fin rays 6; total caudal-fin rays 26; lateral
line with 28 (27–29) tubed scales, posteriormost large,
posterior to hypural crease on caudal fin; median predorsal
scales 13 (12–13); scales above lateral line to origin of dorsal
fin 4 (4–5); scales above anal-fin origin to lateral line 9/10 (9–
10); circumpeduncular scales 16 (one paratype with 18); gill
rakers 18 (13–18); branchiostegals 6.
Vertebrae 9þ16; first vertebral centrum about half width of
subsequent centra, bearing relatively short neural spine;
single supraneural, 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; 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, 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 associated segmented ray and insert 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; pu2 haemal
spine autogenous; 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 with-
out apparent hypurapophysis (Fig. 2B).
Body elongate, greatest depth 3.7 (3.7–4.6) in SL, width 2.3
(1.9–2.4) in greatest depth; head pointed, length 3.1 (3.1–3.5)
in SL; dorsal profile of head slightly convex; snout moder-
ately long, 3.1 (2.7–3.3) in HL; caudal peduncle short and
narrow, least depth 2.8 (2.5–3.0) in HL, its length 2.7 (2.5–
3.4) in HL. Mouth small, terminal, upper jaw nearly reaching
Fig. 2. Radiographs of selected species of Pseudojuloides. Images not to scale. (A) Pseudojuloides atavai, male paratype, AMS I.21716-001, 89.1
mm SL; (B) Pseudojuloides pluto, new species, male paratype, BPBM 40391, 88.9 mm SL; (C) Pseudojuloides proserpina, new species, male
holotype, BPBM 39270, 91.0 mm SL. Radiographs by A. Hay and S. E. Reader.
Tea et al.—Two new species of Pseudojuloides 683
Fig. 3. Pseudojuloides pluto, new species: (A–C) freshly euthanized males showing various degrees of intensity in black body coloration. (A) BPBM
40392, male holotype, 78.0 mm SL, Wake Island, Micronesia; (B) BPBM 40391, male paratype, 88.4 mm SL, Maug Islands, Northern Mariana
Archipelago; (C) BPBM 40391, male paratype, 88.9 mm SL, Maug Islands, Northern Mariana Archipelago; (D) male holotype in preservation; (E)
freshly euthanized female paratype, BPBM 40391, 57.6 mm SL, Maug Islands, Northern Mariana Archipelago; (F) female paratype in preservation,
BPBM 41375, 65.3 mm SL, Wake Island, Micronesia. Photographs by B. D. Greene (A–C, E) and Y. K. Tea (D, F).
Fig. 4. Underwater photographs of Pseudojuloides pluto, new species, from Ogasawara Islands, Japan, at 20 m. (A) Male, approximately 100 mm TL;
(B) female, approximately 80 mm TL; (C) male, approximately 100 mm TL; (D) female, approximately 90 mm TL. Photographs by S. Kobayashi.
684 Copeia 108, No. 3, 2020
a point vertically below anterior nostril. A pair of large,
moderately projecting, and slightly recurved canine 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 7 (6–7; upper jaw) and 8
(8–9; lower jaw) irregularly placed chisel-like incisiform teeth
along each side of jaws.
Preopercular margin smooth, upper part free to between
level of corner of mouth and lower orbit; lower margin free to
a vertical at anterior edge of orbit. Nostrils small, in front of
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 horizontal
peduncular portion; lateral-line scales with a single pore.
Scales cycloid, moderately large except those on breast 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 (3.0–4.1) in HL and 1.2 (1.0–1.8) in longest soft
dorsal-fin ray, third anal-fin spine 4.2 (3.4–4.5) in HL and 1.6
(1.2–1.6) in longest soft anal-fin ray; caudal fin truncate, 1.5
(1.4–1.6) in HL; pectoral fins small, 1.9 (1.7–2.3) in HL; origin
of pelvic fins below lower base of pectorals, their length 2.1
(2.0–2.3) in HL.
Coloration of males in life.—Based on color photographs of
specimens when freshly dead, and live individuals in the
field and aquaria (Figs. 3A–C, 4A, C, 5): head and upper body
olive green, fading to whitish green ventrally; head with a
pair of stripes originating from upper lip; first stripe pale pink
to lilac, sometimes sinuous and weakly bifurcating, tracing
lower edge of orbit to upper and lower edges of operculum;
second stripe pinkish red, continuing dorsally to upper edge
of orbit, extending past nape and base of dorsal fin to upper
edge of caudal peduncle; iris dark olive green, with bright
yellow ring around pupil; scales on lower part of abdomen
edged faintly in lilac forming a honeycomb pattern; anterior
abdominal region behind pectoral and pelvic fins with
narrow oblique lilac to orangey pink bar, about width of
orbit, from dorsal-fin origin to pelvic fin; this ground
coloration often entirely obscured in black save for light
markings on abdomen, head, and dorsal-fin base; dorsal fin
fuliginous to black, interspinous membrane between anterior
two to three spines of dorsal fin with an irregular black spot,
sometimes edged in metallic blue; caudal fin dusky with inky
blue wash, outer edge with a hyaline wedge about one-third
width of caudal fin; inner edge of hyaline portion metallic
blue; anal fin dusky and unmarked, edged narrowly in
metallic blue; pelvic fins grayish hyaline; pectoral fins
hyaline.
Coloration of females in life.—Based on color photographs of
specimens when freshly dead, and live individuals in the
field and aquaria (Figs. 3E, 4B, D, 5): upper and lower body
delimited by a brilliant blue stripe originating from upper lip,
through orbit just beneath lower edge of pupil, continuing
dorsolaterally towards upper edge of caudal peduncle; upper
body brick red to orange-brown, gradually darkening to black
more extensively at interface with blue stripe; lower body,
lower head, and caudal peduncle white to cream, sometimes
pinkish towards caudal-fin base; iris yellowish-orange, lower
one-third below lower edge of pupil white; pupil encircled in
bright orange; dorsal fin orangey hyaline, interspinous
membrane between anterior two to three spines of dorsal
fin with a black spot, sometimes edged in metallic blue and
yellow; caudal fin orangey hyaline; anal fin orangey hyaline,
edged narrowly in metallic blue; pelvic fins hyaline; pectoral
fins pinkish hyaline.
Coloration in alcohol.—(Fig. 3D, F) Males entirely dark tan,
dorsal one-third of body darker with a brownish lateral stripe
from behind upper edge of operculum to upper edge of
caudal peduncle; dorsal, caudal, and anal fins fuliginous;
black spot on anterior dorsal fin remains; pelvic and pectoral
fins translucent; females similar to color in life, except body
uniformly tan and blue body stripe darkening to brown;
dorsal, caudal, anal, and pelvic fins hyaline; black spot on
anterior dorsal fin remains; pectoral fins translucent.
Habitat and distribution.—Pseudojuloides pluto is described on
the basis of nine specimens from the Maug Islands in the
Northern Mariana Island chain, and seven specimens from
Wake Island, northeastern Micronesia. Within the Northern
Mariana Island chain, it has been reported from Tinian and
Guam (Springer, 1982; Kato, 2016). Photographs in the
Image Database of Fishes, Kanagawa Prefectural Museum of
Fig. 5. Head detail of Pseudojuloides pluto, new species, female on left, male with muted coloration on right. Note anterior dorsal fin spot in both
sexes and reduced facial reticulations in the male. Aquarium specimens from the Maug Islands, Northern Mariana Archipelago. Photographs by B. D.
Greene.
Tea et al.—Two new species of Pseudojuloides 685
Natural History (KPM) and records from the literature
indicate that the species also occurs in Saipan (KPM-NR
28259, KPM-NR 28285; Myers and Donaldson, 2003; Kuiter,
2010) and the Ogasawara Islands, Japan (KPM-NR 22317; Fig.
4). It inhabits patch reefs on sandy rubble at depths ranging
from 3–30 m.
Etymology.—The specific epithet refers to Pluto, god and king
of the underworld in Roman mythology, in reference to the
nefarious and sinister appearance of the males. To be treated as
a noun in apposition. The contrasting beauty of the females is
reminiscent of Narcissus flowers, a sacred symbol of Pluto that
grew along the River Styx at the boundary of the underworld.
Pseudojuloides proserpina, new species
urn:lsid:zoobank.org:act:6C1FEDC1-AFE2-40CD-B7ED-
1FB33481CD90
Pomegranate Pencil Wrasse
Figure 6, Table 1
Holotype.—BPBM 39270, 91.0 mm SL, northwest face of Fatu
Hiva, Marquesas, French Polynesia, 25 m, rubble terrace
adjacent to drop off, spear, J. L. Earle, 23 August 2003 (Fig. 6).
Diagnosis.—The following combination of characters distin-
guishes Pseudojuloides proserpina from congeners: body
elongate (greatest body depth 28% SL); dorsal-fin rays
IX,11; anal-fin rays III,12; pectoral-fin rays 13; tubed lateral-
line scales 28; interspinous membrane between anterior two
to three spines of dorsal fin with a black spot; head yellow-
green and extensively reticulate in males; abdominal region
behind pectoral and pelvic fins orangey pink, scales edged in
lavender forming a honeycomb pattern; breast and posterior
body inky black.
Description.—Dorsal-fin rays IX,11, all segmented rays
branched; anal-fin rays III,12, all segmented rays branched;
pectoral-fin rays 13, upper two unbranched; pelvic-fin rays
I,5; principal caudal-fin rays 7þ7; upper and lowermost
unbranched; upper procurrent caudal-fin rays 6; lower
procurrent caudal-fin rays 6; total caudal-fin rays 26; lateral
line with 28 (damaged on right side from spear wound) tubed
scales, posteriormost large, posterior to hypural crease on
caudal fin; median predorsal scales 11; scales above lateral
line to origin of dorsal fin 4; scales above anal-fin origin to
lateral line 9; circumpeduncular scales 16; gill rakers 16;
branchiostegals 6.
Fig. 6. (A) Freshly euthanized and (B) preserved specimen of Pseudojuloides proserpina, new species, BPBM 39270, 91.0 mm SL, male holotype,
Fatu Hiva, Marquesas. Photographs by L. A. Rocha and Y. K. Tea, respectively.
686 Copeia 108, No. 3, 2020
Vertebrae 9þ16; first vertebral centrum about half width of
subsequent centra, bearing relatively short neural spine;
single supraneural, 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; 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, 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 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; pu2 haemal
spine autogenous; 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 with-
out apparent hypurapophysis (Fig. 2C).
Body elongate, greatest depth 3.6 in SL, width 1.1 in
greatest depth; head pointed, length 3.4 in SL; dorsal profile
of head slightly convex; snout moderately long, 3.0 in HL;
caudal peduncle short and narrow, least depth 2.5 in HL, its
length 3.0 in HL. Mouth small, terminal, upper jaw nearly
reaching a point vertically below anterior nostril. A pair of
large, moderately projecting, and slightly recurved canine
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 (upper
jaw) and 6 (lower jaw) irregularly placed chisel-like incisiform
teeth along each side of jaws.
Preopercular margin smooth, upper part free to between
level of corner of mouth and lower orbit; lower margin free to
a vertical at anterior edge of orbit. Nostrils small, in front of
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 horizontal
peduncular portion; lateral-line scales with a single pore.
Scales cycloid, moderately large except those on breast 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 3.4 in HL and 1.2 in longest soft dorsal ray, third
anal-fin spine 4.0 in HL and 1.3 in longest soft anal ray;
caudal fin truncate 1.6 in HL; pectoral fins small, 1.8 in HL;
origin of pelvic fins below lower base of pectorals, their
length 2.1 in HL.
Coloration of males in life.—Based on color photographs of the
holotype when freshly dead (Fig. 6A): head dusky yellow-
green; upper lip with a pair of sinuous maroon stripes to
anterior edge of orbit, becoming increasingly reticulate and
anastomose past orbit toward edge of opercle; stripe on lower
edge of orbit bifurcating, with extensions to lower edge of
maxilla and preopercle; base of dorsal fin with a pink stripe;
iris yellow-green, with bright yellow ring around pupil; scales
on body uniformly purple black except abdominal region
from behind pectoral fin to anal-fin origin, where scales are
bright orange pink, edged narrowly in lilac forming a
honeycomb pattern; dorsal fin yellow, interspinous mem-
brane between anterior first to second spines of dorsal fin
with a black spot; caudal fin purple black on upper and lower
lobes, central rays and distal edge translucent orange, giving
appearance of a forked caudal fin; anal fin dusky yellow,
narrowly orange on distal edge; pelvic fins translucent
yellow; pectoral fins hyaline orange.
Coloration of females in life.—Unknown (but see Comparisons
section below).
Coloration in alcohol.—(Fig. 6B) Head and body uniformly
tan; orange abdominal region becomes pale tan; dorsal fin
translucent, anterior black spot remains; caudal fin fuligi-
nous, anal fin translucent; pectoral fin translucent.
Habitat and distribution.—Pseudojuloides proserpina is known
only from the male holotype collected from Fatu Hiva,
Marquesan Islands, along a rubble terrace adjacent to a drop
off at 25 m (Fig. 7). It shares this distribution with the
endemic P. pyrius. Given the high levels of endemism in the
Marquesas (13.7%; third only to the Hawaiian and Easter
Islands; Delrieu-Trottin et al., 2015), it is likely that P.
proserpina is also endemic to the region.
Etymology.—The specific epithet refers to Proserpina, goddess
and queen of the underworld in Roman mythology, in
reference to both its haunting coloration and close relation-
ship to P. pluto. To be treated as a noun in apposition. The
honeycomb pattern behind the pectoral fin bears a resem-
blance to the seed-studded interior of the pomegranate fruit,
a sacred symbol that features prominently in the myth of
Proserpina and her abduction by Pluto into the underworld.
Remarks.—Although known from only the male holotype, P.
proserpina differs from P. atavai and P. pluto on the basis of
meristic counts, morphometrics, and coloration pattern (see
below). Despite extensive sampling effort, the species has
only been observed once, at 25 m in Fatu Hiva. It is likely
that the species frequents much deeper reefs below depths of
30 m. This could account for its presumed rarity, as
mesophotic reefs in the Marquesan Islands have been
comparatively less well explored. Pseudojuloides proserpina
occurs at depths close to the maximum recorded limit of P.
atavai and P. pluto, and has not been reported above 25 m
where both species commonly occur.
It is well understood that coloration plays an important
role in labrid evolution, being some of the fastest-evolving
phenotypic characters for many species. Presumably, these
are a result of intense sexual selection, where males develop
intricate, yet consistent coloration patterns to attract females
of the correct species. This is well documented for several
genera of sexually dimorphic labrids, including Pseudojuloides
(Victor and Randall, 2014). Unlike P. atavai, the facial
markings of P. proserpina are yellow green with deep purple
reticulations. It is possible that these coloration patterns have
evolved to accommodate a preference for deeper waters.
Whether these differences in coloration pattern are linked to
its ecological preference and mate choice selection requires
further study.
Comparisons.—Both Pseudojuloides pluto and P. proserpina
most closely resemble P. atavai (Fig. 1), sharing similarities
in meristics, morphometrics, and coloration details. Molec-
ular analysis of mitochondrial COI supports a close relation-
Tea et al.—Two new species of Pseudojuloides 687
ship between P. pluto and P. atavai (6.8% uncorrected pairwise
distance). We were unable to retrieve usable DNA sequence
data from the preserved holotype of P. proserpina.
The three species (hereafter referred to as the Pseudojuloides
atavai complex) are united in having a combination of
coloration details in both sexes that are not found in other
species of Pseudojuloides. Unlike the females of other
congeneric species, which are uniformly or suffused green
to red, the females of P. pluto and P. atavai are distinctly
bicolored. While the female form of P. proserpina is not
known, we believe the species is closely allied to P. atavai
based on coloration details of the males. Males of all three
species share the following combination of coloration details:
first two to three interspinous membrane of the dorsal fin
with a black spot; head with a network of reticulate stripes
(reduced in P. pluto); dorsal-fin base with a pink stripe;
abdominal region behind pectoral and pelvic fins pale lilac to
orangey pink (width of pale region dependent on species)
with a crosshatch or honeycomb pattern; and extensive black
coloration over at least posterior half of body. Differences in
live coloration detail are summarized in Table 2.
Aside from details in live coloration, P. pluto differs from P.
atavai in having: a shorter pelvic-fin spine (8.3–10.4% SL vs.
9.5–11.7% SL); a smaller pectoral fin (14.3–18.1% SL vs.
17.7–19.1% SL); more median predorsal scales (12–13 vs. 10–
12); and a different longest dorsal-fin ray (9
th
or 10
th
vs. 1
st
).
Pseudojuloides proserpina differs from P. atavai in having: a
deeper body (28.0% vs. 22.6–25.1% SL for greatest body
depth; 24.9% vs. 21.2–22.0% SL for depth at dorsal-fin
origin); a shorter snout (9.8% vs. 10.8–11.5% SL); a longer
dorsal fin (59.7% vs. 55.1–58.4% SL); a shorter third anal-fin
spine (7.4% vs. 8.5–10.8% SL); and a shorter pelvic-fin spine
(8.8% vs. 9.5–11.7% SL). Pseudojuloides proserpina differs from
P. pluto in having: fewer median predorsal scales (11 vs. 12–
13) and a different longest dorsal-fin ray (1
st
vs. 9
th
or 10
th
).
Given that Pseudojuloides proserpina is known only from the
holotype, differences in morphometric and meristic values
should be taken as tentative pending availability of more
specimens. Nonetheless, both new species are sufficiently
distinct from each other and from P. atavai to warrant species
recognition, and although molecular sequence data for P.
proserpina is unavailable, we believe, based on similarities in
coloration details of the males, that the three species
comprise a monophyletic group.
Phylogenetic interpretation.—Previous phylogenetic analyses
using mitochondrial COI show support for three lineages
within the genus Pseudojuloides (Victor, 2017). The first is
comprised of two species complexes—the cerasinus complex,
consisting of P. c e r a s i n u s ,P. kaleidos,P. p o l a c k o r u m ,P.
polynesica,P. pyrius,P. splendens,P. xanthomos; and the severnsi
complex, consisting of P. edwardi,P. labyrinthus,P. mesostig-
Fig. 7. Distribution records for selected species of Pseudojuloides based on examined specimens and literature records: purple circles,
Pseudojuloides pluto, new species; orange hexagon, Pseudojuloides proserpina, new species; pink stars, Pseuojuloides atavai. Type localities
represented by outlined symbols.
688 Copeia 108, No. 3, 2020
ma,P. severnsi, and P. zeus. In Victor’s (2017) molecular
phylogeny, these complexes share a sister relationship, and
together account for over 70 percent of the species in the
genus. Members from this lineage are typically coral reef
associated, are fairly small, and have females that are
unmarked with an orange to pink suffusion throughout the
body. The 13 species are distributed across the Indian and
Pacific Oceans, from the east African coast to the Polynesian
Islands.
The remaining lineages are very divergent and are
represented in Victor’s (2017) molecular phylogeny by the
species P. atavai and P. elongatus. Of the two, P. elongatus differ
among congeners in having one additional dorsal-fin ray (12
vs. 11), one fewer pectoral-fin ray (12 vs. 13), no median
predorsal scales, and an additional posterior canine on the
upper jaw. Tea et al. (2020, in this volume) revised the
taxonomy of P. elongatus, described two additional species,
and defined the group as the Pseudojuloides elongatus
complex. Members from this complex are anti-equatorial,
preferring macroalgal reefs in subtropical to warm temperate
latitudes.
We expand Victor’s (2017) phylogenetic study with the
inclusion of P. crux,P. paradiseus, and P. pluto. Molecular
phylogenetic analysis inferred using maximum likelihood
and Bayesian inference yielded tree topologies similar to that
of Victor (2017), but with P. atavai having a sister-group
relationship with P. pluto and with P. elongatus forming a
clade with P. crux and P. paradiseus (Fig. 8). Both lineages are
reciprocally monophyletic, but because sequence data for P.
proserpina is unavailable, we consider the sister relationship
of P. atavai and P. pluto as provisional, and the relationship of
the P. atavai complex as unresolved. In both analyses, the
Pseudojuloides elongatus complex is retrieved as the sister
lineage to all remaining Pseudojuloides. We note, however,
that several nodes were recovered with low support,
particularly along the backbone of the tree. Owing to the
limitations of a single locus dataset, we are at this time
unable to confidently resolve the phylogenetic relationships
within the genus. In addition to P. proserpina, molecular
sequence data for P. argyreogaster and P. e r y t h r o p s are
unavailable. We expect a number of these relationships to
change with the use of multi-locus datasets with greater
taxon coverage.
DATA ACCESSIBILITY
Alignment and newick files of phylogenetic trees in this
study are available at https://www.copeiajournal.org/ ci-19-
327. GenBank and BOLD accession numbers are included in
the supplemental appendix (GenBank accession numbers for
the new species Pseudojuloides pluto are MN638808–11).
ACKNOWLEDGMENTS
The holotype and the paratypes of Pseudojuloides pluto were
collected by J. L. Laughlin. We thank A. Hay and A.
Suzumoto for providing curatorial assistance and access to
material at the Australian Museum and Bishop Museum,
respectively. We thank J. T. Williams, E. Fleishauer, R.
Winterbottom, and R. S. Smith and G. Edgar of Reef Life
Survey for providing excellent photographs of Pseudojuloides
atavai. We also thank S. Kobayashi for the excellent in situ
photographs of Pseudojuloides pluto, and L. A. Rocha for
photographs of the holotype of P. proserpina. X-radiographs
were provided by A. Hay and S. Reader. Additional material of
Pseudojuloides pluto used for molecular analyses was collected
in accordance with the Wake Island Natural Resource Permit.
We thank D. P. Brown and Capt. W. Miles for issuing
collection permits, and R. L. Pyle and S. Moncrief for
assisting.
Table 2. Summary of live coloration characters for selected species of Pseudojuloides. Unless otherwise stated, characters apply only to males in
terminal phase.
Pseudojuloides atavai
Pseudojuloides pluto,
new species
Pseudojuloides proserpina,
new species
Facial reticulations Lilac to mauve; extensive,
sinuous, and anastomosed,
from upper lip to preopercle
and opercle
Lilac to light pink; minimal, one or
two sinuous stripes, from upper lip
past lower edge of orbit to upper
edge of opercle
Maroon; extensive, sinuous,
and anastomosed, from
upper lip to preopercle and
opercle
Abdominal region behind
pectoral and pelvic fins
Orange to orangey pink with
yellow crosshatch markings
Narrow lilac to orangey pink oblique
bar, about width of orbit; lilac
honeycomb markings usually
obscured by black body coloration
Orange with lilac honeycomb
markings
Degree of black body markings Present only on posterior half
of body, past anal-fin origin
Present everywhere, often marring
entire body and head save for
stripes on head and narrow bar on
abdomen
Present on breast and posterior
half of body past anal-fin
origin
Dorsal fin Yellow Fuliginous Yellow
Caudal fin Dusky, hyaline centrally and on
outer portion, appearing
forked
Dusky, hyaline on outer edge, about
one-third width of caudal fin,
appearing truncate
Dusky, hyaline centrally and on
outer portion, appearing
forked
Anal fin Dusky yellow Fuliginous Dusky yellow
Coloration of females Orange upper body darkening
abruptly to black narrowly at
interface with blue lateral
stripe
Orange upper body darkening
gradually to black more extensively
at interface with blue lateral stripe
Unknown
Tea et al.—Two new species of Pseudojuloides 689
Fig. 8. Maximum-likelihood and Bayesian phylogenetic relationships of Pseudojuloides inferred using mitochondrial COI. Support values at nodes
correspond to posterior probability and maximum-likelihood bootstrap percentages, respectively. The new species, Pseudojuloides pluto (purple), is
at present most closely related to P. atavai (pink). The Pseudojuloides cerasinus, severnsi, and elongatus complexes are in green, blue, and yellow,
respectively. See Data Accessibility for tree files.
690 Copeia 108, No. 3, 2020
LITERATURE CITED
Bouckaert, R., J. Heled, D. K¨
uhnert, T. Vaughan, C.-H. Wu,
D. Xie, M. A. Suchard, A. Rambaut, and A. J. Drum-
mond. 2014. BEAST 2: a software platform for Bayesian
evolutionary analysis. PLoS Computational Biology 10:
e1003537.
Chang, C. H., K. T. Shao, H. Y. Lin, Y. C. Chiu, M. Y. Lee,
and P. L. Lin. 2017. DNA barcodes of the native ray-finned
fishes in Taiwan. Molecular Ecology Resources 17:796–805.
Delrieu-Trottin, E., J. T. Williams, P. Bacchet, M. Kulbicki,
J. Mourier, R. Galzin, T. Lison de Loma, G. Mou-Tham,
G. Siu, and S. Planes. 2015. Shore fishes of the Marquesas
Islands, an updated checklist with new records and new
percentage of endemic species. Check List 11:1753.
Edgar, R. C. 2004. MUSCLE: multiple sequence alignment
with high accuracy and high throughput. Nucleic Acids
Research 32:1792–1797.
Gill, A. C., Y. K. Tea, and H. Senou. 2016. Plectranthias
takasei, a new species of anthiadine fish from southern
Japan (Teleostei: Serranidae). Zootaxa 4205:349–356.
Johnson, G. D., and C. Patterson. 2001. The intermuscular
system of acanthomorph fishes: a commentary. American
Museum Novitates 3312:1–24.
Kato, S. 2016. Wrasses and Parrotfishes Illustrated. Seibundo
Shinkosha, Tokyo.
Kuiter, R. H. 2010. Labridae Fishes: Wrasses. Aquatic Photo-
graphics, Seaford, Australia.
Lanfear, R., P. B. Frandsen, A. M. Wright, T. Senfeld, and B.
Calcott. 2016. PartitionFinder 2: new methods for select-
ing partitioned models of evolution for molecular and
morphological phylogenetic analyses. Molecular Biology
and Evolution 34:772–773.
Lobel, P. S., and L. K. Lobel. 2004. Annotated checklist of
the fishes of Wake Atoll. Pacific Science 58:65–90.
Myers, R. F. 1988. An annotated checklist of fishes of the
Mariana Islands. Micronesica 21:115–180.
Myers, R. F., and T. J. Donaldson. 2003. The fishes of the
Mariana Islands. Micronesica 3536:594–648.
Patterson, C., and G. D. Johnson. 1995. The intermuscular
bones and ligaments of teleostean fishes. Smithsonian
Contributions to Zoology 559:1–83.
Rambaut, A., A. Drummond, D. Xie, G. Baele, and M. A.
Suchard. 2018. Posterior summarization in Bayesian
phylogenetics using Tracer 1.7. Systematic Biology 67:
901–904.
Randall, J. E. 1999. Report on fish collections from the
Pitcairn Islands. Atoll Research Bulletin 461:1–53.
Randall, J. E., and H. A. Randall. 1981. A revision of the
labrid fish genus Pseudojuloides, with descriptions of five
new species. Pacific Science 35:51–74.
Ronquist, F., M. Teslenko, P. van der Mark, D. L. Ayres, A.
Darling, S. Hohna, B. Larget, L. Liu, M. A. Suchard, and
J. P. Huelsenbeck. 2012. MrBayes 3.2: efficient Bayesian
phylogenetic inference and model choice across a large
model space. Systematic Biology 61:539–542.
Russell, B. C. 1988. Revision of the labrid fish genus
Pseudolabrus and allied genera. Records of the Australian
Museum Supplement 9:1–72 (plus addendum after plates),
pls 1–4.
Sabaj, M. H. 2019. Standard symbolic codes for institutional
resource collections in herpetology and ichthyology: An
Online Reference. Version 7.1 (21 March 2019). Electron-
ically accessible at http://www.asih.org, American Society
of Ichthyologists and Herpetologists, Washington, D.C.
(accessed 8 July 2019).
Springer, V. G. 1982. Pacific Plate Biogeography, with special
reference to shorefishes. Smithsonian Contributions to
Zoology 367:1–162.
Stamatakis, A. 2014. RAxML version 8: a tool for phyloge-
netic analysis and post-analysis of large phylogenies.
Bioinformatics 30:1312–1313.
Tea, Y. K., A. C. Gill, and H. Senou. 2020. Two new species
of Pseudojuloides from Western Australia and southern
Japan, with a redescription of Pseudojuloides elongatus
(Teleostei: Labridae). Copeia 108.
Victor, B. C. 2017. Review of the Indo-Pacific Pseudojuloides
cerasinus species complex with a description of two new
species (Teleostei: Labridae). Journal of the Ocean Science
Foundation 29:11–31.
Victor, B. C., and J. E. Randall. 2014. Pseudojuloides edwardi,
n. sp. (Perciformes: Labridae): an example of evolution of
male-display phenotype outpacing divergence in mito-
chondrial genotype. Journal of the Ocean Science Foun-
dation 11:1–12.
Tea et al.—Two new species of Pseudojuloides 691
