Cirrhilabrus finifenmaa (Teleostei, Labridae), a new species of fairy wrasse from the Maldives, with comments on the taxonomic identity of C. rubrisquamis and C. wakanda

Abstract

Cirrhilabrus rubrisquamis is redescribed on the basis of the juvenile holotype and compared to known species of Cirrhilabrus . Examination of material from the Maldives identified as C. rubrisquamis reveal differences from the holotype collected from the Chagos Archipelago. Consequently, the Maldivian specimens are herein described as Cirrhilabrus finifenmaa sp. nov. , on the basis of the holotype and twelve paratypes. The new species differs from all congeners in having: males with anterior third to half of body bright magenta, peach to orange-pink posteriorly; lateral line with 22–26 pored scales (16–18 in the dorso-anterior series, 6–8 in the posterior peduncular series); tenth to eleventh dorsal-fin spine longest (14.0–15.5% SL); scales on the opercle, chest, isthmus, and anterior third of the body with a dark purple-red central region (purple in alcohol), the markings joining appearing crosshatched; dorsal, caudal, anal, and pelvic-fin rays purple in alcohol. Meristic details and coloration patterns of C. rubrisquamis are very similar to C. wakanda from Tanzania, Africa, although synonymy of both species cannot be determined without additional material from Chagos. This potential synonymy is briefly discussed; however, until such material becomes available, the taxonomic statuses of C. wakanda and C. rubrisquamis are here provisionally regarded as valid.

Full text

Cirrhilabrus finifenmaa (Teleostei, Labridae),
a new species of fairy wrasse from the Maldives,
with comments on the taxonomic identity
of C. rubrisquamis and C. wakanda
Yi-Kai Tea1,2, Ahmed Najeeb3, Joseph Rowlett4, Luiz A. Rocha5
1School of Life and Environmental Sciences, University of Sydney, Sydney, Australia 2Ichthyology, Australian
Museum Research Institute, 1 William Street, Sydney, New South Wales 2010, Australia 3Maldives Marine
Research Institute, Ministry of Fisheries Marine Resources and Agriculture, Malé 20025, Maldives 4Field
Museum of Natural History, Chicago, Illinois, 60605, USA 5Department of ichthyology, California Academy
of Sciences, San Francisco, California 94118, USA
Corresponding author: Yi-Kai Tea (yi-kai.tea@sydney.edu.au)
Academic editor: Bruno Melo|Received 18 November 2021|Accepted 14 February 2022|Published 8 March 2022
http://zoobank.org/6404BD85-20F7-4017-ADA8-113C2DBF97EB
Citation: Tea Y-K, Najeeb A, Rowlett J, Rocha LA (2022) Cirrhilabrus nifenmaa (Teleostei, Labridae), a new species
of fairy wrasse from the Maldives, with comments on the taxonomic identity of C. rubrisquamis and C. wakanda.
ZooKeys 1088: 65–80. https://doi.org/10.3897/zookeys.1088.78139
Abstract
Cirrhilabrus rubrisquamis is redescribed on the basis of the juvenile holotype and compared to known spe-
cies of Cirrhilabrus. Examination of material from the Maldives identied as C. rubrisquamis reveal dier-
ences from the holotype collected from the Chagos Archipelago. Consequently, the Maldivian specimens
are herein described as Cirrhilabrus nifenmaa sp. nov., on the basis of the holotype and twelve paratypes.
e new species diers from all congeners in having: males with anterior third to half of body bright ma-
genta, peach to orange-pink posteriorly; lateral line with 22–26 pored scales (16–18 in the dorso-anterior
series, 6–8 in the posterior peduncular series); tenth to eleventh dorsal-n spine longest (14.0–15.5% SL);
scales on the opercle, chest, isthmus, and anterior third of the body with a dark purple-red central region
(purple in alcohol), the markings joining appearing crosshatched; dorsal, caudal, anal, and pelvic-n rays
purple in alcohol. Meristic details and coloration patterns of C. rubrisquamis are very similar to C. wakanda
from Tanzania, Africa, although synonymy of both species cannot be determined without additional mate-
rial from Chagos. is potential synonymy is briey discussed; however, until such material becomes avail-
able, the taxonomic statuses of C. wakanda and C. rubrisquamis are here provisionally regarded as valid.
Keywords
Coral reefs, deep reefs, Indian Ocean, mesophotic, reef sh
ZooKeys 1088: 65–80 (2022)
doi: 10.3897/zookeys.1088.78139
https://zookeys.pensoft.net
Copyright Yi-Kai Tea et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Introduction
Randall and Emery (1983) described Cirrhilabrus rubrisquamis from the 40.6 mm
standard length (SL) holotype collected at 41–48 m depth in Isla Fouquet, Peros
Banhos Atoll, Chagos Archipelago. eir description was based on a single juvenile
specimen, and so appropriate details regarding adult coloration and characters were
unavailable. In his subsequent review of Cirrhilabrus from the Western Indian
Ocean, Randall (1995) rediagnosed C. rubrisquamis with additional material from
Vilingili Island, North Malé Atoll, Maldives, thus expanding its distribution, and
stating that “it is satisfying to be able to diagnose and illustrate the adult female
and male of this colorful species”. Although Randall’s specimens consisted mostly of
mature individuals of both sexes, it also included a juvenile that diered in coloration
pattern from the similarly-sized holotype from Chagos, thus raising the question
as to whether the specimens from the Maldives represent a dierent species. Re-
examination of the aforementioned Maldivian specimens indicate that they indeed
represent a dierent species from the nominal Chagossian C. rubrisquamis. To resolve
this taxonomic conundrum, the Maldivian specimens are examined and compared
with C. rubrisquamis sensu stricto and related congeners, and here described as the
new species Cirrhilabrus nifenmaa. e holotype of C. rubrisquamis is redescribed
and compared to related Cirrhilabrus. It is shown to be a valid species of the genus
and a possible senior synonym of C. wakanda. We briey comment on the taxonomic
status of both species, but provisionally regard both as valid pending additional
material of adult C. rubrisquamis from Chagos.
Materials and methods
Methods for counting and measuring mostly follow Randall and Masuda (1991).
Gill rakers are presented as upper (epibranchial) + lower (ceratobranchial); the an-
gle raker is included in the second count. Counts of lateral-line scales are given in
two parts, the dorso-anterior series and the midlateral peduncular series. e lat-
ter series consists of a larger pored scale overlapping the caudal-n base, which we
include in the count. In the description that follows, data are presented rst for the
holotype, followed by the range of minimum–maximum values of the paratypes in
parentheses where dierent. Where counts were recorded bilaterally, both counts
are given and separated from each other by a slash; the rst count presented is the
left count. Morphometric values are expressed as percentage of standard length
(Table 1). Osteological details were taken from radiographs of the holotype and
paratypes. Institutional codes follow Sabaj (2020) and are as follows: Australian
Museum, Sydney (AMS); Bernice Pauahi Bishop Museum (BPBM); Royal On-
tario Museum (ROM); California Academy of Sciences (CAS); Zoological Refer-
ence Collection of the Lee Kong Chian Natural History Museum at the National
University of Singapore (ZRC).

A new species of fairy wrasse from the Maldives 67
Taxonomy
Cirrhilabrus rubrisquamis Randall & Emery, 1983
Rosy-scaled Fairy Wrasse
Fig. 1, Table 1
Cirrhilabrus rubrisquamis Randall & Emery, 1983: 21; g. 1 (description); Winter-
bottom et al. 1989: 53; pl VII-E (checklist, shes of the Chagos Archipelago);
Winterbottom and Anderson 1997: 15 (revised checklist, shes of the Chagos
Archipelago)
Holotype. Cirrhilabrus rubrisquamis: ROM 35932, 40.6 mm SL, juvenile, Isla Fou-
quet, Peros Banhos Atoll, Chagos Archipelago (5°21'3"S, 71°48'57"E).
Diagnosis. Cirrhilabrus rubrisquamis shares similar meristic characters to other
members of this genus, in particular C. wakanda (the potential synonymy of both spe-
cies is discussed below). However, the holotype is readily distinguished from related
congeners in having the following combination of characters: lateral line with 21 or 22
pored scales (15 or 16 in the dorso-anterior series, six in the posterior peduncular se-
ries); gill rakers 16; caudal n round with blue and yellow vermiculation in life; dorsal
two-thirds of body with purple scales arranged in a chain-link pattern in life.
Description. Dorsal-n rays XI,9; all soft rays branched except rst; anal-n rays III,9;
all soft rays branched except rst; last dorsal and anal-n ray branched to base; pectoral-n
rays 15/15, upper two unbranched; pelvic-n rays I,5; principal caudal-n rays 7+6, up-
permost and lowermost unbranched; upper procurrent caudal-n rays 5, lower procurrent
caudal-n rays 5; lateral line interrupted, with dorso-anterior series of pored scales 16/15
and midlateral posterior peduncular series 6/6; rst pored scale on posterior peduncular
series often pitted; last pored scale on posterior peduncular series enlarged and overlapping
hypural crease; scales above lateral line to origin of dorsal n 2; scales below lateral line to
origin of anal n 6; median predorsal scales 5; median prepelvic scales 5; rows of scales on
cheek 2; circumpeduncular scales 16; gill rakers 5+11 = 16; pseudobranchial lament
count not made, owing to small size of specimen; vertebrae 9 + 16; epineurals 12.
Body moderately elongate and compressed, depth 3.5 in SL, width 2.2 in depth;
head length (HL) 2.9 in SL; snout pointed, its length 3.6 in HL; orbit diameter 3.5
in HL; depth of caudal peduncle 2.1 in HL. Mouth small, terminal, and oblique,
with maxilla almost reaching vertical at front edge of orbit; dentition typical of genus
with three pairs of canine teeth present anteriorly at side of upper jaw, rst forward-
projecting, next two strongly recurved and outcurved, third longest; an irregular row of
very small conical teeth medial to upper canines; lower jaw with a single stout pair of
canines anteriorly which protrude obliquely outward and are slightly lateral to medial
pair of upper jaw; no teeth on roof of mouth.
Posterior margin of preopercle with 29/27 very ne serrations; margins of posterior
and ventral edges of preopercle free to about level of middle pupil. Anterior nostril in
short membranous tube, located nearer to orbit than snout tip; posterior nostril larger,

Yi-Kai Tea et al. / ZooKeys 1088: 65–80 (2022)
68
Table 1. Proportional measurements for holotype of Cirrhilabrus rubrisquamis, and type series of C. nifenmaa sp. nov., and C. wakanda, expressed as percentage
of the standard length. Data for C. wakanda (n = 5) summarized from Tea et al. (2019).
Cirrhilabrus
rubrisquamis
Cirrhilabrus nifenmaa sp. nov. Cirrhilabrus wakanda
ROM 35932 BPBM
33094
BPBM 41385 (formerly BPBM 33094) ZRC
62259
ZRC
62249
ZRC
61604
AMS I.50058-001 CAS-ICH 247311
(exAMS I.50058-001)
CAS 246395– CAS246399
Holotype Holotype Paratypes Holotype and four
paratypes
Sex Juvenile Male Male Male Female Female Juvenile Male Male Female Male Male Female Female 2 Males and three females
Standard length (mm) 40.6 69.2 70.5 57.6 52.9 52.1 35.8 76.7 54.2 47.9 69.1 59.7 54.0 57.2 54.3–70.3
Body depth 28.8 31.6 30.2 32.3 32.3 29.8 35.8 31.3 28.2 29.0 31.4 31.2 28.3 29.4 29.8–31.9
Body width 13.3 13.9 11.8 13.2 14.2 13.2 12.6 14.6 13.1 12.1 17.2 16.1 15.2 15.7 11.8–14.5
Head length 35.0 31.6 30.4 31.3 32.1 32.4 33.2 31.0 33.4 35.9 33.9 33.5 32.6 33.0 27.7–31.2
Snout length 9.6 9.5 8.8 9.0 8.1 8.1 9.5 8.5 8.5 10.2 9.1 8.4 8.1 8.4 7.4–8.9
Orbit diameter 10.1 8.8 9.5 9.2 10.2 10.2 11.7 9.6 10.3 11.1 9.0 9.7 10.0 9.1 6.6–9.0
Interorbital width 7.4 9.1 8.2 9.5 9.8 9.4 9.5 8.9 9.2 9.0 9.3 10.2 9.4 10.0 7.7–9.6
Upper jaw length 6.4 7.4 5.8 8.7 7.0 7.9 8.1 9.3 5.7 7.3 5.8 5.7 4.6 4.2 6.5–8.3
Caudal-peduncle depth 16.5 14.5 15.5 15.1 15.9 15.9 15.4 14.3 15.9 16.3 17.7 16.9 14.8 15.4 14.8–16.5
Caudal-peduncle length 13.5 15.8 17.9 16.5 15.9 15.2 10.1 18.9 11.8 14.8 15.1 15.1 13.1 14.3 12.8–16.5
Predorsal length 36.9 31.2 31.9 33.3 31.6 35.9 34.4 33.4 34.9 38.4 33.6 34.0 33.0 34.1 31.7–33.8
Preanal length 65.3 61.4 61.0 60.1 67.3 61.8 65.6 55.7 58.1 61.8 57.6 59.6 61.7 61.0 58.5–61.4
Prepelvic length 40.9 35.7 35.7 34.7 34.4 34.9 38.3 33.5 34.3 38.6 35.0 34.5 34.6 33.9 31.5–36.5
Dorsal-n base 57.4 60.4 57.7 60.8 60.7 58.0 57.5 61.3 58.5 61.4 60.0 61.0 57.6 61.0 55.3–63.2
First dorsal spine 9.1 7.9 6.8 8.0 7.4 7.5 7.5 6.8 7.0 7.3 5.6 6.7 7.4 6.8 5.2–6.5
Longest dorsal spine 15.0 15.5 15.0 15.1 15.5 15.2 14.0 15.6 15.3 15.9 10.6 14.2 15.6 14.3 11.9–14.3
Longest dorsal ray 17.2 19.1 20.1 18.1 18.1 17.7 damaged 21.4 15.3 20.0 21.7 17.1 18.1 18.7 16.7–19.0
Anal-n base 22.4 28.0 24.5 24.8 25.7 22.8 25.1 26.6 26.0 24.0 28.5 25.6 27.0 26.9 25.3–27.6
First anal spine 8.9 6.4 7.4 6.4 6.4 5.8 5.9 6.0 6.1 6.5 6.4 4.9 8.3 6.6 5.2–6.4
Second anal spine 13.5 9.7 10.9 10.8 11.0 11.3 10.6 11.0 12.4 10.2 11.1 9.7 11.5 10.1 9.1–10.1
ird anal spine 14.3 11.6 11.2 11.8 12.1 13.2 11.7 12.3 12.5 10.2 13.0 11.7 12.2 11.4 10.5–11.4
Longest anal ray 18.0 20.2 19.7 17.0 17.0 16.9 14.8 23.3 20.5 21.1 21.7 20.1 16.9 18.4 14.5–17.9
Caudal-n length 29.6 27.5 29.8 25.9 28.0 28.8 32.1 30.0 33.6 30.7 28.5 29.8 29.6 32.9 25.4–31.6
Pectoral-n length 19.7 20.2 20.0 21.0 21.4 20.9 19.3 23.5 19.7 24.0 6.4 21.8 20.7 21.5 18.3–21.8
Pelvic-spine length 14.8 11.3 11.5 12.2 12.7 13.4 13.4 11.5 12.0 12.9 11.1 11.4 11.9 11.0 11.0–12.1
Pelvic-n length 19.2 22.7 17.2 18.8 20.8 19.2 20.4 22.9 18.6 18.4 13.0 16.8 17.8 21.7 15.5–18.8

A new species of fairy wrasse from the Maldives 69
roughly ovoid to rectangular, located just medial and anterior to upper edge of eye.
Scales cycloid; head scaled except snout and interorbital space; six large scales on opercle;
a broad naked zone on membranous edge of preopercle; a row of large, elongate, pointed
scales along base of dorsal n, one per element, scales progressively shorter posteriorly on
soft portion of n; anal n with a similar basal row of scales; last pored scale of lateral line
(posterior to hypural plate) enlarged and pointed; one scale above and below last pored
scale also enlarged; pectoral ns naked except for a few small scales at extreme base; a
single large scale at base of each pelvic n, about three-fourths length of pelvic spine.
Origin of dorsal n above second or third lateral-line scale, predorsal length 2.7 in
SL; rst 1–5 dorsal-n spines progressively longer, sixth to tenth subequal, eleventh long-
est, 2.3 in HL; interspinous membranes of dorsal n in males extend beyond dorsal-n
spines, with each membrane extending in a pointed cirri beyond spine; eighth dorsal-n
soft ray longest, 2.0 in HL, remaining rays progressively shorter; origin of anal n be-
low base of tenth dorsal-n spine; third anal-n spine longest, 2.4 in HL; interspinous
membranes of anal n extended as on dorsal n; anal-n soft rays relatively uniform in
length, eighth longest, 1.9 in HL; dorsal and anal-n rays just reaching caudal-n base;
caudal n rounded; pectoral ns short, reaching vertical between bases of fourth or fth
dorsal-n spines, longest ray 1.8 in HL; origin of pelvic ns below lower base of pectoral
ns; pelvic ns short, not reaching past anal n origin, longest ray 1.8 in HL.
Coloration of holotype in life. Based on color photograph of holotype when
fresh (Fig. 1): head yellow, brownish posteriorly; lower part of head whitish to pale
pink; preopercle prominently purple on outer edge; iris yellow; body orangey pink,
fading to whitish pink ventrally; body with a network of dark purple scales arranged in
a chain link pattern from just after dorsal n origin to edge of caudal peduncle, absent
from lower third of body; dorsal n hyaline, yellow on distal half; posterior dorsal n
yellowish hyaline with metallic blue spots; caudal n bluish hyaline with yellow and
blue vermiculation, often broken in spots; anal n similar to dorsal n; pelvic ns hya-
line; pectoral n base with a purple band; pectoral ns pinkish hyaline.
Figure 1. Cirrhilabrus rubrisquamis, ROM 35932, 40.6 mm SL, juvenile holotype, Isla Fouquet, Peros
Banhos Atoll, Chagos Archipelago. Photograph by AR Emery and R Winterbottom.

Yi-Kai Tea et al. / ZooKeys 1088: 65–80 (2022)
70
Coloration of holotype in alcohol. Uniformly pale tan, median and paired ns
translucent hyaline. No evidence of purple scale markings, likely due to immaturity
and/or loss of coloration over time.
Cirrhilabrus nifenmaa sp. nov.
http://zoobank.org/E6D891D1-FAB0-45B6-BE5A-D4123BF85A13
Rose-Veiled Fairy Wrasse
Figs 2–4; Table 1
Cirrhilabrus rubrisquamis (non Randall & Emery, 1983): Randall and Anderson,
1993: pl. 6C (checklist, underwater photograph from Maldives); Randall 1995:
gs 5–7 (preserved specimens, BPBM 33094); Hawkins et al. 2000: 85 (IUCN
assessment); Allen et al. 2008: (in part [Maldivian distribution], checklist of valid
species of Cirrhilabrus); Kuiter 2010: 135 (color photographs A–C; aquarium
specimens from Maldives); Allen et al. 2015: (in part [Maldivian distribution],
checklist of valid species of Cirrhilabrus); Hawkins et al. 2016: g. 1 (uorescence
spectrometry of a male specimen); Tea and Gill 2017: g. 8H (color photograph
of an aquarium specimen from Maldives); Tea et al. 2018: g. 10 (preserved speci-
men, BPBM 33094); Tea et al. 2019: g. 5D (color photograph of an aquarium
specimen from Maldives); Tea et al. 2021a (included as part of a phylogenomic
study of the genus).
Holotype. BPBM 33094, 69.2 mm SL, male, east coast of Vilingili Island, North
Malé Atoll, Maldives, 52 m, collected by John E. Randall with quinaldine and spear,
23–25 March 1988 (Fig. 2A).
Paratypes. BPBM 41385 (formerly BPBM 33094) (5), 35.8–70.5 mm SL, 1 juve-
nile, 2 females, 2 males, collected with male holotype (Fig. 2C, E); AMS I.50058-001
(3), 54.0–69.1 mm SL, 2 males, 1 female, Hulhumalé Island, North Malé Atoll, Mal-
dives (4°14'6.66"N, 73°33'15.47"E), 35–40 m, collected by Ibrahim Rasheed with
hand nets, 15 August 2021 (Fig. 2B1–B2, D1–D2, F1–F2); CAS-ICH 247311 (ex
AMS I.50058-001), 57.2 mm SL, female, collected with AMS I.50058-001; ZRC
62259, 76.7 mm SL, male, aquarium specimen from Maldives (Figs 2G, 3); ZRC
62249, 54.2 mm SL, male, aquarium specimen from Maldives; ZRC 61604, 47.9 mm
SL, female, aquarium specimen from Maldives.
Diagnosis. A species of Cirrhilabrus distinguished from congeners based on the
following combination of characters: males with anterior third to half of body bright
magenta, peach to orange-pink posteriorly; lateral line with 22–26 pored scales (16–18
in the dorso-anterior series, 6–8 in the posterior peduncular series); tenth to eleventh
dorsal-n spine longest (14.0–15.5% SL); scales on the opercle, chest, isthmus, and
anterior third of the body with a dark purple-red central region, the markings joining
to form a crosshatched appearance (purple in alcohol); dorsal, caudal, anal, and pelvic-
n rays purple in alcohol.

A new species of fairy wrasse from the Maldives 71
Description. Dorsal-n rays XI,9; all soft rays branched except rst; anal-n rays
III,9; all soft rays branched except rst; last dorsal and anal-n ray branched to base;
pectoral-n rays 15 (right side removed in all three ZRC paratypes), upper two un-
branched; pelvic-n rays I,5; principal caudal-n rays 7+6, uppermost and lowermost
unbranched; upper procurrent caudal-n rays 6 (6–7), lower procurrent caudal-n
rays 6 (5–6); lateral line interrupted, with dorso-anterior series of pored scales 17/17
(16–18) and midlateral posterior peduncular series 8/7 (6–8); rst pored scale on pos-
terior peduncular series often pitted; last pored scale on posterior peduncular series
enlarged and overlapping hypural crease; scales above lateral line to origin of dorsal n
2; scales below lateral line to origin of anal n 6 (6–7); median predorsal scales 5 (4–5);
Figure 2. Cirrhilabrus nifenmaa, sp. nov., not to scale A male holotype, BPBM 33094, 69.2 mm SL,
Vilingili Island, North Malé Atoll, Maldives B1, B2 male paratype, AMS I.50058-001, 69.1 mm SL,
Hulhumalé Island, North Malé Atoll, Maldives, in life and in preservation respectively C young male
paratype, BPBM 41385 (formerly BPBM 33094), 57.6 mm SL, same data as holotype D1, D2 young
male paratype, AMS I.50058-001, 59.7 mm SL, Hulhumalé Island, North Malé Atoll, Maldives, in life
and in preservation respectively E juvenile paratype, BPBM 41385 (formerly BPBM 33094), 35.8 mm
SL, same data as holotype F1, F2 female paratype, AMS I.50058-001, 54.0 mm SL, Hulhumalé Island,
North Malé Atoll, Maldives, in life and in preservation respectively G male paratype in life, ZRC 62259,
76.7 mm SL, aquarium specimen from Maldives H male in nuptial colors, aquarium specimen from
Maldives, specimen not retained. Photographs by the late JE Randall (A, C , E), AN (B1–B2, D1–D2,
F1–F2), and YKT (G, H).

Yi-Kai Tea et al. / ZooKeys 1088: 65–80 (2022)
72
Figure 3. Cirrhilabrus nifenmaa sp. nov., male paratype, ZRC 62259, 76.7 mm SL, (A) freshly
euthanized (B) and in preservation. Note (C) purple scale markings forming a crosshatched pattern and
(D)purple n rays in preservation. Photographs by YKT.
median prepelvic scales 6 (5–6); rows of scales on cheek 2; circumpeduncular scales 14
(14–16); gill rakers 5 (5–6) + 11 (9–12) = 16 (14–18); pseudobranchial laments 11
(10–11); vertebrae 9 + 16; epineurals 13.
Body moderately elongate and compressed, depth 3.2 (2.8–3.5) in SL, width 2.3
(1.8–2.8) in depth; head length (HL) 3.2 (2.8–3.3) in SL; snout pointed, its length
3.3 (3.3–4.0) in HL; orbit diameter 3.6 (2.8–3.8) in HL; depth of caudal peduncle
2.2 (1.9–2.2) in HL. Mouth small, terminal, and oblique, with maxilla almost reach-
ing vertical at front edge of orbit; dentition typical of genus with three pairs of canine
teeth present anteriorly at side of upper jaw, rst forward-projecting, next two strongly

A new species of fairy wrasse from the Maldives 73
recurved and outcurved, third longest; an irregular row of very small conical teeth
medial to upper canines; lower jaw with a single stout pair of canines anteriorly which
protrude obliquely outward and are slightly lateral to medial pair of upper jaw; no
teeth on roof of mouth.
Posterior margin of preopercle with 35/36 (27–39) very ne serrations; margins of
posterior and ventral edges of preopercle free to about level of middle pupil. Anterior
nostril in short membranous tube, located nearer to orbit than snout tip; posterior
nostril larger, roughly ovoid to rectangular, located just medial and anterior to upper
edge of eye. Scales cycloid; head scaled except snout and interorbital space; 6 (6–7)
large scales on opercle; a broad naked zone on membranous edge of preopercle; a row
of large, elongate, pointed scales along base of dorsal n, one per element, scales pro-
gressively shorter posteriorly on soft portion of n; anal n with a similar basal row of
scales; last pored scale of lateral line (posterior to hypural plate) enlarged and pointed;
one scale above and below last pored scale also enlarged; pectoral ns naked except for
a few small scales at extreme base; a single large scale at base of each pelvic n, about
three-fourths length of pelvic spine.
Origin of dorsal n above second or third lateral-line scale, predorsal length 3.2
(2.6–3.2) in SL; rst 1–5 dorsal-n spines progressively longer, sixth to ninth subequal,
tenth to eleventh longest, 2.0 (2.0–2.4) in HL; interspinous membranes of dorsal n
in males extend beyond dorsal-n spines, with each membrane extending in a pointed
cirri beyond spine; eighth to ninth dorsal-n soft ray longest (three paratypes with
second dorsal-n soft ray longest), 1.6 (1.5–2.3) in HL, remaining rays progressively
shorter; origin of anal n below base of 10th dorsal-n spine; third anal-n spine long-
est, 2.7 (2.4–3.5) in HL; interspinous membranes of anal n extended as on dorsal n;
anal-n soft rays relatively uniform in length, seventh to ninth longest, 1.6 (1.5–2.2)
in HL; dorsal and anal-n rays just reaching past caudal-n base; caudal n rounded;
pectoral ns short, reaching vertical between bases of 6th or seventh dorsal-n spines,
longest ray 1.6 (1.3–1.7) in HL; origin of pelvic ns below lower base of pectoral ns;
pelvic ns short, not reaching past anal n origin, longest ray 1.4 (1.4–2.0) in HL.
Coloration of males in life. Based on color photograph of holotype and paratypes
when fresh, and live specimens photographed in the eld and aquaria (Figs 2A, B1,
C, D1, G, H, 3A, 4): head magenta to orange-red, red stripe present from mid-upper
lip to mid-upper edge of orbit, continuing to anterior half of dorsal n base; second
stripe of similar color present from lower edge of maxilla to mid-lower edge of orbit;
nape to predorsal region with four to ve thin white stripes; preopercle prominently
purple on outer edge; iris bright yellow with red ring around pupil; anterior third of
body magenta to orange-red pink, remaining two thirds peach to orange-pink; scales
on opercle, chest, isthmus and anterior third of body with dark purple-red central
region, markings joining to form a crosshatched appearance; dorsal n orange, some-
times ush with pink, becoming progressively hyaline posteriorly; outer edge of dorsal
n metallic blue; segmented dorsal-n rays deep purple; basal two-thirds of caudal n
magenta to rose-red, remaining third metallic blue; caudal-n rays deep purple; anal
n similar to dorsal n; pelvic ns translucent orange, rays deep purple.

Yi-Kai Tea et al. / ZooKeys 1088: 65–80 (2022)
74
Coloration of females and juveniles in life. Based on color photographs of holo-
type and paratypes when freshly dead, and live specimens photographed in the eld
and aquaria (Fig. 2E, F1): similar to male, except distinction between magenta head
and anterior body from paler posterior two-thirds less prominent, sometimes suused;
coloration on median ns less obvious, appearing hyaline or pale orange-pink; juve-
niles similar to females, except uniformly suused-pink; median ns pinkish hyaline.
Coloration in alcohol. Similar to coloration in life, except head and body light
tan to cream; edge of preopercle purple; dark purple-red scale markings now purple;
dentary, Angulo-articular, and bony edge of preopercle purple; segmented rays of dor-
sal, anal, caudal, and pelvic ns purple (Figs 2B2, D2, F2, 3B–D).
Etymology. e epithet is from the Dhivehi "nifenmaa", meaning rose, alluding
to the live coloration of this species. e pink rose yatoshi nifenmaa (Rosa spp.) is
also the national ower of the Maldives. To be treated as a noun in apposition. e
common name is given after the facial patterns of the species.
Habitat and distribution. Cirrhilabrus nifenmaa is presently known from Mal-
dives and Sri Lanka, at depths ranging from 40–70 m (Fig. 4). e species belongs to
the C. jordani species complex, a group of deep-water fairy wrasses found mostly in
mesophotic coral ecosystems. It is likely that this species occurs in greater depths. Like
other species of Cirrhilabrus, C. nifenmaa frequents rubble bottoms scattered with
loose coral cover.
Remarks. Mitochondrial COI and 16S sequences for C. nifenmaa are publicly
available on GenBank (accession numbers MH780161 and MH780159 respectively;
previously identied and listed as C. rubrisquamis). We exercise caution in using other
sequences presently identied as belonging to C. rubrisquamis, as these are likely to be
C. nifenmaa.
Material examined. Cirrhilabrus rubrisquamis: ROM 35932, 40.6 mm SL,
juvenile holotype, Isla Fouquet, Peros Banhos Atoll, Chagos Archipelago. Cirrhilabrus
nifenmaa: BPBM 33094, 69.2 mm SL, male holotype, east coast of Vilingili Island,
North Malé Atoll, Maldives, 52 m, collected by John E. Randall with quinaldine and
spear, 23–25 March 1988; BPBM 41385 (formerly BPBM 33094) (5), 35.8–70.5mm
SL, 1 juvenile, 2 females, 2 males, paratypes, collected with male holotype (Fig. 2C, E);
AMS I.50058-001 (3), 54.0–69.1 mm SL, 2 males, 1 female, paratypes, Hulhumalé
Island, North Malé Atoll, Maldives, 35–40 m, collected by Ibrahim Rasheed with
hand nets, 15 August 2021; CAS-ICH 247311 (ex AMS I.50058-001), 57.2mm SL,
female, collected with AMS I.50058-001; ZRC 62259, 76.7 mm SL, male paratype,
aquarium specimen from Maldives (Figs 2G, 3); ZRC 62249, 54.2 mm SL, male
paratype, aquarium specimen from Maldives; ZRC 61604, 47.9 mm SL, female
paratype, aquarium specimen from Maldives; Cirrhilabrus wakanda: CAS 246395,
70.3 mm SL, male holotype, east coast of Zanzibar, Tanzania, Africa, 75 m, collected
by H.T. Pinheiro, B. Shepherd, and L.A. Rocha, 14 December 2018; CAS 246396,
56.8 mm SL, female paratype, east coast of Zanzibar, Tanzania, Africa, 70 m, 07
December 2018; CAS 246397, 61.3 mm SL, male paratype, same data as holotype;
CAS 246398, 57.4 mm SL, female paratype, same data as holotype; CAS 246399,
54.3 mm SL, female paratype, same data as holotype.

A new species of fairy wrasse from the Maldives 75
Taxonomic status of Cirrhilabrus rubrisquamis and comparisons to
related species
In their description of Cirrhilabrus rubrisquamis, Randall and Emery (1983) could not
reliably determine the sex of the holotype due to the absence of mature gonads. ey
note the presence of a blunt genital papilla (which we conrm), and while suggestive of
the holotype being female, is dicult to conrm due to the small size of the specimen.
Distinction between juveniles and females for many species of Cirrhilabrus on the basis
of external characters is often dicult, with few reliable coloration characters separating
these life stages. Depending on the species, Cirrhilabrus may possess one or more of the
following coloration characters that feature prominently in the juvenile phase, but later
disappearing or attenuating with maturity: presence of several ne, thread-like stripes
interspersed with white spots on the upper half of the body; presence of a large white
spot on the tip of the snout (Kuiter 2010); presence of a black caudal peduncular spot.
e color photograph of the holotype in Randall and Emery (1983; also included here in
Fig.1) shows the presence of ne, thread-like stripes interspersed with white spots situ-
ated dorsally, below the dorsal n origin. is combination of typical juvenile coloration
pattern and the absence of mature gonads strongly suggests that the holotype is a juvenile.
Cirrhilabrus rubrisquamis is most closely related to C. apterygia from Western
Australia, C. blatteus from the Red Sea, the newly described C. nifenmaa from the
Maldives, C. sanguineus from Mauritius, and C. wakanda from East Africa. e six
species form a clade of Indian Ocean species that belong to a larger group of fairy
Figure 4. Cirrhilabrus nifenmaa sp. nov., underwater photograph from Rasdhoo Atoll, Maldives, 60 m.
Photograph by LAR.

Yi-Kai Tea et al. / ZooKeys 1088: 65–80 (2022)
76
wrasses known as the C. jordani species complex, with species distributed across the
Indo-Pacic (Tea et al. 2021b). Of the aforementioned species, the newly described
C. nifenmaa is most distinct in having a rounded caudal n (vs. lanceolate in
C.apterygia, C. blatteus, C. sanguineus, and C. wakanda [terminal males not known in
C.rubrisquamis; but see below]), in having purple-red crosshatch markings on its opercle,
chest, isthmus, and anterior body, and in having unmarked scales on the posterior
two-thirds of its body. On the basis of meristic, morphometric, and morphological
details, C. nifenmaa further diers from: C. apterygia in having pelvic ns (absent in
C. apterygia); C.blatteus in having a longer snout (8.1–9.5% SL vs. 7.5–8.1% SL) and
a longer preanal length (60.1–67.3% SL vs. 52.1–62.5% SL); C.sanguineus in having a
longer snout (8.1–9.5% SL vs. 7.7–8.7% SL); and C. wakanda in having a larger orbit
(8.8–11.7% SL vs. 6.6–9.0% SL) and a taller dorsal n (eight to ninth soft dorsal-n
ray longest, 14.0–15.5% SL vs. sixth longest, 11.9–14.3% SL).
Due to the lack of adequate material and muddled taxonomy of C. rubrisquamis,
appropriate comparisons to related species of Cirrhilabrus is dicult. In particular, open
questions remain regarding the taxonomic status of C. rubrisquamis and C.wakanda. Tea et
al. (2019) described C. wakanda based on material collected from Zanzibar, Tanzania. ey
compared their new species with related Cirrhilabrus from the Indian Ocean, including
C. rubrisquamis. Although their comparative data for C.rubrisquamis included published
data of the holotype from Chagos, it also included published data from Randall’s (1995)
Maldivian specimens (= C. nifenmaa). While the taxonomy of C. rubrisquamis sensu lato is
here resolved with the description of C.nifenmaa from Maldives, comparative data taken
from the C. rubrisquamis holotype and Randall’s Maldivian material (=C.nifenmaa) in
the description of C. wakanda resulted in a potentially new problem.
Both C. rubrisquamis and C. wakanda are most similar on the basis of coloration
patterns, in particular the presence of purple chain-link markings across the dorsal two-
thirds of the body. Both species also share similar morphometric details (Table1); how-
ever, this is not uncommon in closely related species of Cirrhilabrus, where identication
is most reliable through comparison of terminal males (Tea et al. 2021a). Comparison of
live coloration details between the smallest (54.3 mm SL) paratype of C. wakanda from
Tea et al. (2019) and the 40.6 mm SL holotype of C. rubrisquamis reveal several minor
coloration dierences, such as C. rubrisquamis having a yellow dorsal n (vs. fuchsia in
C. wakanda), and a hyaline caudal n with yellow and blue vermiculation (vs. caudal
n with a magenta chevron marking in C. wakanda). e holotype of C. rubrisquamis
further diers from C. wakanda in having fewer pored lateral line scales (21–22 vs.
24–28). However, since C. rubrisquamis is known only from the juvenile holotype, it is
unclear whether these dierences represent distinct characters or ontogenetic variation.
Recent underwater ROV explorations by the University of Plymouth Research Ex-
pedition to Egmont Atoll and Sandes Seamount in the Chagos Archipelago recovered
in situ footage of fairy wrasses resembling C. rubrisquamis taken at depths between
60–70 m (Fig. 5A–C). Given the close proximity to Peros Banhos Atoll (also in the
Chagos Archipelago), we strongly suspect these to be the nominal C. rubrisquamis.
Examination of live coloration details of putative females (Fig. 5B) taken from the
video footage agree well with the juvenile holotype of C. rubrisquamis and females of

A new species of fairy wrasse from the Maldives 77
C. wakanda (Fig. 5D), particularly in having similar purple chain-link markings. e
putative males (Fig. 5A, C) are similar in appearance to the East African C. wakanda
(Fig. 5E, F), diering only slightly in having a yellow dorsal n (Fig. 5C; vs. fuchsia in
C. wakanda) and in lacking the yellowish saddle in the middle of the body (Fig. 5E,
F). Given the similar live coloration of C. rubrisquamis and C. wakanda, it is possible
that the two nominal species are conspecic. Observable dierences (or potential lack
thereof) in live coloration details however may be attributable to dierences in size of
the specimens, poor video resolution, and dierences in underwater light properties.
Figure 5. Underwater photographs of Cirrhilabrus from the Western Indian Ocean A harem of putative
Cirrhilabrus rubrisquamis. Male depicted in circular inset, females by white arrowheads B putative female
C. rubrisquamis C putative terminal male C. rubrisquamis; Photographs taken from video footage provided
by K Howell, N Foster, and C Diaz from the University of Plymouth Research Expedition to Egmont
Atoll and Sandes Seamount in the Chagos Archipelago, 60–70 m D female C. wakanda, underwater
photograph from Zanzibar, Tanzania, Africa, 75 m E male C. wakanda, underwater photograph from
Zanzibar, Tanzania, Africa, 75 m F terminal male C. wakanda, underwater photograph from Moyette, o
the coast of Mozambique, 100 m. Note fuchsia dorsal n in all individuals and pale yellowish saddle in
males. Photographs by LR (D, E) and P Plantard (F).

Yi-Kai Tea et al. / ZooKeys 1088: 65–80 (2022)
78
e distribution of C. wakanda in East Africa and C. rubrisquamis in the Chagos
Archipelago presents a unique pattern of biogeography shared by few congeneric
Cirrhilabrus. Within the Indian Ocean, there appears to be faunal distinction between
the Chagos Archipelago from the rest of the Western and Eastern Indian Ocean in some,
but not all taxa. is biogeographic connectivity has been reported for corals (Obura
2012), Acanthaster sea stars (Haszprunar et al. 2017), and coconut crabs (Sheppard
et al. 2013), and amongst shes, at least 54 species are known to have a Western
Indian Ocean distribution encompassing the Chagos Archipelago (Winterbottom
and Anderson 1997). Pertinent examples of allopatric species conforming to this
biogeographic pattern include the labrids Halichoeres iridis (East Africa to Chagos) and
H. leucoxanthus (Maldives to Java), and two apparent species of Dascyllus, currently
confused as D. carneus (see Gill and Kemp 2002). Marine shore sh endemism is
particularly low in the Chagos Archipelago (Winterbottom et al. 1989), but with
few notable examples (e.g., Amphiprion chagosensis). Without additional material of
C.rubrisquamis of appropriate sizes from the type locality however, we are unable to
reconcile the taxonomic status of both species, and whether or not C. rubrisquamis
and C. wakanda represent closely related allopatric species, or whether they represent
a single widespread conspecic taxon. Until such material from Chagos becomes
available, we refrain from placing C. wakanda in the synonymy of C. rubrisquamis,
and provisionally regard both species as valid.
Acknowledgements
We thank Amanda Hay and Kerryn Parkinson (AMS), Kelvin Lim and Heok-Hui Tan
(ZRC), Dave Catania (CAS), and Arnold Suzomoto, Calder Atta, Loreen O’Hara,
Rirchard Pyle, and Brian D. Greene (BPBM) for curatorial assistance, loan of speci-
mens, and provision of registration numbers. Marg Zur, Erling Holm, and Rick Win-
terbottom (ROM) assisted in providing data, photographs, and x-radiographs of the
Cirrhilabrus rubrisquamis holotype. We thank Amanda Hay and Kerryn Parkinson
for x-radiographs. Kerry Howell, Nicola Foster, and Clara Diaz provided in situ video
footage of Cirrhilabrus taken from the University of Plymouth Research Expedition to
Egmont Atoll and Sandes Seamount in the Chagos Archipelago. Patrick Plantard pro-
vided excellent underwater photographs used in this manuscript. We thank Mohamed
Ashwag Ismail of Maldives Marine Research Institute (MMRI) for his assistance in
specimen preparation and photography. We thank Ibrahim Rasheed of Sub Tropical
Exotic Paragon (STEP) for collecting the AMS paratypes used in this study. Speci-
mens were collected by STEP, under authorization of MMRI, holding permit numbers
A2021024. Financial support for L. Rocha was provided by donors to the California
Academy of Sciences’ Hope for Reefs Initiative and a Rolex Award for Enterprises.
Mark Westneat and one anonymous reviewer provided comments that improved the
quality of this manuscript.

A new species of fairy wrasse from the Maldives 79
References
Allen GR, Drew J, Barber P (2008) Cirrhilabrus beauperryi, a new wrasse (Pisces: Labridae)
from Melanesia. Aqua (Neu-Isenburg, Germany) 14: 129–140.
Allen GR, Erdmann MV, Dailami M (2015) Cirrhilabrus marinda, a new species of wrasse
(Pisces: Labridae) from eastern Indonesia, Papua New Guinea, and Vanuatu. Journal of the
Ocean Science Foundation 15: 1–13.
Gerlach T, eobald J, Hart NS, Collin SP, Michiels NK (2016) Fluorescence characterization
and visual ecology of pseudocheilinid wrasses. Frontiers in Zoology 13(1): e13. https://doi.
org/10.1186/s12983-016-0145-1
Gill AC, Kemp JM (2002) Widespread Indo-Pacic shore-sh species: A challenge for taxono-
mists, biogeographers, ecologists, and shery and conservation managers. Environmental
Biology of Fishes 65(2): 165–174. https://doi.org/10.1023/A:1020044616889
Haszprunar G, Vogler C, Wörheide G (2017) Persistent gaps of knowledge for naming and
distinguishing multiple species of crown-of-thorns-seastar in the Acanthaster planci species
complex. Diversity (Basel) 9(2): e22. https://doi.org/10.3390/d9020022
Hawkins JP, Roberts CM, Clark V (2000) e threatened status of restricted-range coral reef sh
species. Animal Conservation 3(1): 81–88. https://doi.org/10.1111/j.1469-1795.2000.
tb00089.x
Kuiter RH (2010) Labridae Fishes: Wrasses. First edition. Aquatic Photographics, Seaford, Aus-
tralia, 398 pp.
Obura D (2012) e diversity and biogeography of Western Indian Ocean reef-building corals.
PLoS ONE 7(9): e45013. https://doi.org/10.1371/journal.pone.0045013
Randall JE (1995) A review of the wrasses of the genus Cirrhilabrus (Perciformes: Labridae)
from the Western Indian Ocean. Revue Francaise d’Aquariologie. Herpétologie 22: 19–26.
Randall JE, Anderson RC (1993) Annotated checklist of the epipelagic and shore shes of the
Maldive Islands. J.L.B. Smith Institute of Ichthyology. Ichthyological Bulletin 59: 1–47.
Randall JE, Emery AR (1983) A new labrid sh of the genus Cirrhilabrus from the Chagos
Archipelago, Indian Ocean. Journal of Aquariculture and Aquatic Sciences 3: 21–24.
Randall JE, Masuda H (1991) Two new labrid shes of the genus Cirrhilabrus from Japan.
Revue française d’Aquariologie Herpétologie 18: 53–60.
Sabaj MH (2020) Codes for natural history collections in ichthyology and herpetology. Copeia
108(3): 593–669. https://doi.org/10.1643/ASIHCODONS2020
Sheppard CRC, Bowen BW, Chen AC, Craig MT, Eble J, Fitzsimmons N, Gan CH, Gaither
MR, Gollock M, Keshavmurthy S, Koldewey H, Mortimer JA, Obura D, Pfeier M, Rog-
ers AD, Sheppard ALS, Vogler C, Wöheide G, Yang MC, Yesson C (2013) British Indian
Ocean Territory (the Chagos Archipelago): setting, connections and the marine protected
area. In: Sheppard CRC (Ed.) Coral Reefs of the United Kingdom Overseas Territories.
Springer Nature, Switzerland, 323 pp. https://doi.org/10.1007/978-94-007-5965-7_17
Tea YK, Gill AC (2017) Cirrhilabrus shutmani, a new species of fairy wrasse from the Babuyan
Islands, northern Philippines (Teleostei: Labridae). Zootaxa 4341(1): 77–88. https://doi.
org/10.11646/zootaxa.4341.1.6

Yi-Kai Tea et al. / ZooKeys 1088: 65–80 (2022)
80
Tea YK, Frable BW, Van Der Wal C (2018) Redescription and phylogenetic placement of
Cirrhilabrus sanguineus (Teleostei: Labridae), with rst documentation of the female form.
Zootaxa 4526(3): 358–372. https://doi.org/10.11646/zootaxa.4526.3.5
Tea YK, Pinheiro HT, Shepherd B, Rocha LA (2019) Cirrhilabrus wakanda, a new species of
fairy wrasse from mesophotic ecosystems of Zanzibar, Tanzania, Africa (Teleostei, Labri-
dae). ZooKeys 863: 85–96. https://doi.org/10.3897/zookeys.863.35580
Tea YK, Xu X, DiBattista JD, Lo N, Cowman PF, Ho SYW (2021a) Phylogenomic analysis of
concatenated ultraconserved elements reveals the recent evolutionary radiation of the fairy
wrasses (Teleostei: Labridae: Cirrhilabrus). Systematic Biology 71(1): 1–12. https://doi.
org/10.1093/sysbio/syab012
Tea YK, Allen GR, Goatley CHR, Gill AC, Frable BW (2021b) Redescription of Conniella
apterygia Allen and its reassignment in the genus Cirrhilabrus Temminck and Schlegel (Tel-
eostei: Labridae), with comments on pseudocheiline pelvic morphology. Zootaxa 5061(3):
493–509. https://doi.org/10.11646/zootaxa.5061.3.5
Winterbottom R, Anderson C (1997) A revised checklist of the epipelagic and shore shes
of the Chagos Archipelago, Central Indian Ocean. J.L.B. Smith Institute of Ichthyology.
Ichthyological Bulletin 66: 1–28.
Winterbottom R, Emery AR, Holm E (1989) An annotated checklist of the shes of the Cha-
gos Archipelago Central Indian Ocean. Life Sciences Contributions of the Royal Ontario
Museum 145: 1–226. https://doi.org/10.5962/bhl.title.52237