A New Genus and Species of Pygmy Pipehorse from Taitokerau Northland, Aotearoa New Zealand, with a Redescription of Acentronura Kaup, 1853 and Idiotropiscis Whitley, 1947 (Teleostei, Syngnathidae)

Abstract

Cylix tupareomanaia, new genus and species, is described from three specimens (35.5–55.5 mm SL), collected from rocky reefs at 12–17 m depth from Taitokerau Northland, New Zealand. The new taxon shares morphological synapomorphies with the superficially similar Australian endemic Idiotropiscis and Indo-Pacific Acentronura, including head angled ventrally approximately 25° from the principal body axis, enclosed brood pouch, brood pouch plates, prehensile tail, and absence of caudal fin. Cylix tupareomanaia, new genus and species, however, is distinguishable from all other members of the Syngnathidae by the following combination of bony autapomorphic characters: a cup-like crest present anterodorsally on the supraoccipital; and large conspicuous midventral conical spines on the cleithral symphysis and first trunk ring between the pectoral-fin bases. The new species can be further differentiated by genetic divergence in the mitochondrial COX1 gene from Acentronura breviperula, A. tentaculata, Idiotropiscis australe, and I. lumnitzeri (estimated uncorrected p-distances of 19.5%, 20.4%, 17.9%, and 18.4%, respectively). A phylogenetic hypothesis from the analysis of two nuclear loci, 18S and TMO-4C4, supports the placement of C. tupareomanaia, new genus and species, as the sister taxon to a clade comprising the genera Acentronura and Idiotropiscis. Cylix tupareomanaia, new genus and species, represents the eighth member within the pygmy pipehorse clade to be described from the Indo-Pacific and the first new genus and species of syngnathid to be reported from New Zealand since 1921.

Full text

A New Genus and Species of Pygmy Pipehorse from
Taitokerau Northland, Aotearoa New Zealand, with a
Redescription of Acentronura Kaup, 1853 and
Idiotropiscis Whitley, 1947 (Teleostei, Syngnathidae)
Authors: Short, Graham A., and Trnski, Thomas
Source: Ichthyology & Herpetology, 109(3) : 806-835
Published By: The American Society of Ichthyologists and
Herpetologists
URL: https://doi.org/10.1643/i2020136
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A New Genus and Species of Pygmy Pipehorse from Taitokerau Northland,
Aotearoa New Zealand, with a Redescription of Acentronura Kaup, 1853 and
Idiotropiscis Whitley, 1947 (Teleostei, Syngnathidae)
Graham A. Short
1,2,3
and Thomas Trnski
4
Cylix tupareomanaia, new genus and species, is described from three specimens (35.5–55.5 mm SL), collected from rocky
reefs at 12–17 m depth from Taitokerau Northland, New Zealand. The new taxon shares morphological synapomorphies
with the superficially similar Australian endemic Idiotropiscis and Indo-Pacific Acentronura, including head angled
ventrally approximately 258from the principal body axis, enclosed brood pouch, brood pouch plates, prehensile tail,
and absence of caudal fin. Cylix tupareomanaia, new genus and species, however, is distinguishable from all other
members of the Syngnathidae by the following combination of bony autapomorphic characters: a cup-like crest present
anterodorsally on the supraoccipital; and large conspicuous midventral conical spines on the cleithral symphysis and
first trunk ring between the pectoral-fin bases. The new species can be further differentiated by genetic divergence in
the mitochondrial COX1 gene from Acentronura breviperula,A. tentaculata,Idiotropiscis australe, and I. lumnitzeri
(estimated uncorrected p-distances of 19.5%, 20.4%, 17.9%, and 18.4%, respectively). A phylogenetic hypothesis from
the analysis of two nuclear loci, 18S and TMO-4C4, supports the placement of C. tupareomanaia, new genus and species,
as the sister taxon to a clade comprising the genera Acentronura and Idiotropiscis.Cylix tupareomanaia, new genus and
species, represents the eighth member within the pygmy pipehorse clade to be described from the Indo-Pacific and the
first new genus and species of syngnathid to be reported from New Zealand since 1921.
¯
Anei r¯
a he puninga, he momo manaia iti hou. He uri n¯
o Te Taitamaw¯ahine, n¯
o Te Taitokerau, ar¯
a, n¯
o Aotearoa. Ko Cylix
tupareomanaia t¯
ona ¯
ıngoa. He ¯
ıngoa i tuku mai ¯
aNg
¯
ati Wai. Ka whakaahuatia t¯enei manaia iti mai i ng¯at
¯
ıpako e toru
(35.5–55.5 mitamano te roa paerewa). I kohia mai i te ¯
akau tokatoka o te Te Taitokerau, 12–17 mita te h¯
ohonu. He ¯
ahua
¯
orite ¯
etahi o ng¯
a¯
ahuatanga o t¯
enei r¯
op¯
u hou ki ng¯
a puninga manaia taketake, ko Acentronura,n
¯
oTeP
¯
apaka-a-M¯
aui/
Ahitereiria, me t¯ana whanaunga, ko Acentronura,n
¯
o te rohe kotahi o ¯
Inia me Te Moana-nui-a-Kiwa. Engari, ka taea te
wetewetekia a Cylix tupareomania pun. mo. hou mai i te ¯
ata t¯
ıtiro ki ng¯a ¯
ahuatanga rerek¯eong
¯
ak
¯
oiwi, p¯er¯
a i te putanga
k¯
oiwi mai i te angaanga, me k
¯
ı, he t¯
upare. Ka taea te wetewetekia hoki mai i ng¯
a rerek¯
etanga o ng¯
a ira pata p¯
ungao, ar¯
a
ko te ira COX1 o ng¯
a momo manaia it ko Acentronura breviperula,r
¯
atou ko A. tentacula,ko Idiotropicis australe,ko I.
lumnitzeri. Ka tautoko hoki te m¯atai iranga o ng¯
aw
¯
ahi ira, 18S me TMO-4C4 ki t¯enei whakapae whakapapa pori, hei
r¯
op¯
u whanaunga a C. tupareomanaia, momo hou, ki ng¯
a puninga Acentronura, nga puninga Idiotropicis hoki, ar¯
a, kotahi
te tupuna o ¯
enei manaia katoa. Ko Cylix tupareomania, momo hou, te momo manaia iti tuawaru n¯
o te tupuna manaia
kotahi, n¯
o te rohe kotahi o ¯
Inia me Te Moana-nui-a-Kiwa kua whakaahuatia p¯enei. Me k
¯
ı, ko te tau 1921 k¯e te tau ki mua
ka puta he p¯
urongo p¯enei o t¯etahi puninga hou, momo hou hoki o te wh¯anau manaia.
SYNGNATHIDAE is a large and morphologically dis-
tinctive group of predominantly marine fishes, with 57
genera and 300 described species that include the
seahorses, pipefishes, pygmy pipehorses, and seadragons
(Dawson, 1985; Hamilton et al., 2017; WoRMS Editorial
Board, 2021). Syngnathids primarily inhabit shallow coastal
waters in temperate and tropical oceans, and are character-
ized by male brooding, a unique overall body plan, including
an elongated snout, fused jaw, the absence of pelvic fins,
thick plates of bony armor, and behavioral and morpholog-
ical adaptations for crypsis.
The inshore marine ichthyofauna of New Zealand includes
a small but diverse group of syngnathids represented by ten
species in five genera: Hippocampus,Leptonotus,Lissocampus,
Solegnathus, and Stigmatopora (Dawson, 1980, 1985; Roberts
et al., 2015; Clark et al., 2017). Four species of syngnathids
are endemic to New Zealand, including Lissocampus filum,a
kelp-associated species, Stigmatopora macropterygia, the largest
member of its genus, and two uncommon sand and rubble-
associated species, Leptonotus elevatus and L. norae, that
belong to a Pacific genus with a disjunct distribution with
the only other species, L. blainvilleanus, occurring in South
America. In contrast, syngnathid diversity in nearby Australia
is exceptionally high with 129 species represented in 47
genera, 11 of which are monotypic (Dawson, 1985; Browne
et al., 2008; Baker et al., 2009; Hamilton et al., 2017). All
genera of syngnathids that occur in temperate mainland New
Zealand, with the exception of Leptonotus, occur in southern
Australia (Dawson, 1985). Similarly, New Caledonia, which is
linked to New Zealand by the almost entirely submerged
continental landmass Te Riu-a-M¯
aui Zealandia (Mortimer et
al., 2017), exhibits high syngnathid diversity and endemism
with 41 species in 14 genera (Fricke et al., 2011).
Recent surveys of inshore marine biodiversity at the
offshore subtropical Rangit¯
ahua Kermadec Islands in the
northernmost region of New Zealand produced new records
of Indo-Pacific syngnathids (Francis and Duffy, 2015; Trnski
et al., 2015; Clark et al., 2017). These surveys yielded the
1
Department of Ichthyology, California Academy of Sciences, San Francisco, California 94118; Email: gshort@calacademy.org. Send reprint
requests to this address.
2
Ichthyology, Australian Museum Research Institute, Sydney NSW 2010, Australia.
3
Research Associate, Ichthyology, Burke Museum, Seattle, Washington.
4
Auckland War Memorial Museum T¯
amaki Paenga Hira, Auckland 1142, New Zealand; Email: ttrnski@aucklandmuseum.com.
Submitted: 5 October 2020. Accepted: 15 March 2021. Associate Editor: M. T. Craig.
Ó2021 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/i2020136 Published online: 20 September 2021
Ichthyology & Herpetology 109, No. 3, 2021, 806–835
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pipefish species Cosmocampus howensis,Halicampus boothae,
and the rare southwest Pacific seahorse Hippocampus jugumus,
all of which were originally described based on specimens
collected from Lord Howe Island, Australia (Whitley, 1948;
Allen et al., 1976; Smith et al., 2010). In 2009 and 2014, the
Bay of Islands Coastal Survey Project, coordinated by the
National Institute of Water and Atmospheric Research
(NIWA), conducted extensive marine biodiversity surveys of
shallow water coastal habitats in the Northland region of
New Zealand, including the Bay of Islands and the Cavalli
Islands. These surveys each yielded two individuals of a
diminutive syngnathid (Jones et al., 2009; NIWA, 2014:
https://niwa.co.nz/news/rare-fish-and-new-seafloor-habitats-
found-during-niwa-survey-of-coastal-east-northland) that
were provisionally identified as the Australian pygmy pipe-
horse Idiotropiscis australe based on a superficial similarity to
this species (Stewart, 2015).
Members of the genus Idiotropiscis belong to a grouping of
Indo-Pacific species of pygmy pipehorses comprising the
genera Acentronura and Kyonemichthys, all of which have the
head angled slightly ventrally from the abdominal axis, a
fully enclosed male brood pouch, prehensile tail, and no
caudal fin. The Western Atlantic pygmy pipehorse genus
Amphelikturus appears to have evolved independently in the
Atlantic to the extent that Dawson (1984) suggested all
pygmy pipehorses shared a common ancestor; however,
molecular and morphological differences support the find-
ings of independent evolution of Atlantic and Pacific pygmy
pipehorses (Hamilton et al., 2017). A multi-gene phyloge-
netic hypothesis of relationships of the family Syngnathidae
recovered Acentronura and Idiotropiscis as sister taxa within a
clade of highly morphologically distinct Indo-Pacific pipefish
genera Filicampus,Trachyrhamphus,Haliichthys, and Lisso-
campus (Hamilton et al., 2017). Even though Acentronura and
Idiotropiscis share the synapomorphy of fully enclosed brood
pouch, prehensile tail, and absence of caudal fin with the
seahorse genus Hippocampus, no immediate sister taxon
relationship was supported among these genera (Hamilton
et al., 2017). Instead, a sister-group relationship was
recovered between the Indo-Pacific clade containing Acen-
tronura and Idiotropiscis and a globally distributed and well-
defined clade of seahorses (Hamilton et al., 2017). The three
previously described species of the genus Idiotropiscis,I.
australe (typespeciesofthegenus),I. larsonae,andI.
lumnitzeri, are endemic to the shallow coastal waters of
Australia (Waite and Hale, 1921; Dawson, 1984; Kuiter,
2004). Idiotropiscis australe occurs in South Australia and
southern Western Australia, I. larsonae in northern Western
Australia, and I. lumnitzeri in southern New South Wales.
Although the collected specimens from northern New
Zealand appeared superficially similar in appearance to I.
australe based on shared synapomorphies, such as the head
angled ventrally approximately 258from the principal body
axis, a fully enclosed brood pouch, brood pouch plates,
prehensile tail, and absence of caudal fin (Stewart, 2015),
closer investigation revealed a number of unique morpho-
logical characters present on the head that differed markedly
from I. australe, and with features unknown from any other
species of pygmy pipehorse described previously. On the
basis of these morphological features, molecular sequence
data, and phylogenetic analysis, we consider the species from
Taitokerau, Northland, New Zealand as a new genus and
species of pygmy pipehorse in the family Syngnathidae, and
we describe it herein. Non-type representatives of the
superficially similar Acentronura and Idiotropiscis are rede-
scribed based on a substantially expanded dataset of
morphological characters (compared to the original descrip-
tions), as revealed by micro-computed tomography. Com-
parisons of these morphological characters revealed novel
species-specific neurocranial characters that assisted in the
identification of the generic limits of Acentronura,Idio-
tropiscis, and Cylix, new genus.
MATERIALS AND METHODS
Morphological analyses.—Head and body measurements
reported follow Short et al. (2018) and are expressed as
percent of standard length (SL) or head length (HL).
Osteocranial and postcranial skeletal terminology follows
Leysen (2011). External morphological characters were
documented using a dissecting microscope and analyses of
high-resolution digital images. Morphological characters of
the axial skeleton of type specimens and comparative
material were examined via non-destructive x-ray micro-
computed tomography (lCT) at the Karel F. Liem Bioimaging
Facility (Friday Harbor Laboratories, University of Washing-
ton). Micro-computed tomography (lCT) was performed
using a Bruker Skyscan 1173 scanner (Billerica, MA) with a
1 mm aluminum filter at 60 kV and 110 lA on a 2048 32048
pixel CCD at a resolution of 8.8 lm. The specimens were
placed inside a 50 ml plastic Falcon tube (Corning, NY),
supported by two thin foam pads to prevent movement
during scanning and wrapped in 70% ethanol-infused
cheesecloth to maintain moist conditions and prevent
desiccation. The resulting lCT data were visualized, segment-
ed, and rendered in Horos software (https://horosproject.
org).
Specimens examined are housed in the following museum
collections: Auckland War Memorial Museum (AIM); Austra-
lian Museum, Sydney (AMS); Museum of New Zealand Te
Papa Tongarewa (NMNZ); South Australian Museum, Ade-
laide (SAMA); Western Australian Museum, Perth (WAM);
Museum and Art Gallery of the Northern Territory, Darwin
(NTM). Institutional abbreviations follow Sabaj (2020).
Georeferenced locations for the type specimens of C.
tupareomanaia, new genus and species, were captured on
GPS units and use WGS84 datum.
Molecular analyses.—DNA extraction, primers, PCR condi-
tions, sequence alignment, and analysis of COX1 and
nuclear sequence data generated for this study, as well as
previously published mitochondrial and nuclear sequence
data, were performed following protocols described in
Hamilton et al. (2017). COX1 sequence data for C. tupar-
eomanaia, new genus and species, in this study followed
protocols described in Eme et al. (2020). A partial segment of
mitochondrial cytochrome c oxidase subunit I (COX1) DNA
was sequenced from a tail clipping of a specimen from the
type locality of C. tupareomanaia, new genus and species,
(AIM MA122275) and a specimen of I. australe (SAMA F2657).
COX1 sequence data was compared to the previously
sequenced pygmy pipehorse species A. breviperula,A. tenta-
culata, and I. lumnitzeri (available from Hamilton et al., 2017)
in order to calculate genetic distances (uncorrected p-
distances) in MEGA v. 7.0.26 (Kumar et al., 2018). The
phylogenetic position of the new species was investigated
Short and Trnski—New pygmy pipehorse from New Zealand 807
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among the morphologically similar pygmy pipehorse species
A. breviperula,A. tentaculata, and I. lumnitzeri, and sampled
members of Hippocampus based on the nuclear markers 18s
ribosomal (18S) and anonymous putative titin-like protein
(TMO-4C4), which were derived in this study and from
Hamilton et al. (2017) employing Maximum Likelihood
(ML), Maximum Parsimony (MP), and MrBayes analyses
(Appendix 1). DNA sequences for the two gene markers were
assembled, aligned, and concatenated using Geneious v.
11.1.4 (Biomatters, Ltd., Auckland). The aligned dataset was
imported into MEGA v.7.0.26 (Kumar et al., 2018) for ML and
MP analyses. ML analyses were evaluated using bootstrap-
ping with 1,000 replicates for topological support. The
resulting best scoring ML phylogram was rooted with the
southern Australian trunk-brooder pipefish Heraldia nocturna
and viewed using FIGTREE v.1.3.1 (http://tree.bio.ed.ac.uk/
software/figtree/). MP analyses were conducted with 1,000
bootstrap replicates using a full heuristic search utilizing tree-
bisection and reconnection (TBR) branch swapping. All
characters were weighted equally and unordered. Resulting
equally parsimonious cladograms were rooted with H.
nocturna, summarized using a strict consensus method, and
viewed using FIGTREE v1.3.1. The Bayesian analyses were
performed following protocols described in Hamilton et al.
(2017) using MrBayes v.3.2 (Ronquist et al., 2012) with
settings as follows: GTR substitution model for 18S and TMO-
4C4 with a gamma-distributed rate parameter applied, two
Metropolis-coupled Markov Chain Monte Carlo (MCMC)
runs, and each with four chains. The analyses ran for 10
million generations sampling trees and parameters every
1,000 generations.
Cylix, new genus
urn:lsid:zoobank.org:act:B9144B5E-2220-4C29-ABEA-
D7CCD4288258
Type species.—Cylix tupareomanaia, new species.
Diagnosis.—A genus of the Syngnathidae that shares numer-
ous morphological synapomorphies with Acentronura and
Idiotropiscis, including head angled ventrally approximately
258from the principal body axis, enclosed brood pouch,
brood pouch plates, prehensile tail, and absence of caudal
fin. However, Cylix tupareomanaia, new species, differs from
all other genera by unique anatomical features of the head,
including: a distinct, cup-like crest present anterodorsally on
the supraoccipital; and large and conspicuous medioventral
conical spines on the cleithral symphysis and the first trunk
ring between the pectoral-fin bases. It differs further in
having the following combination of morphological charac-
ters: prominent supraoccipital; continuous cleithrum; prom-
inent supracleithrum; anterior nuchal plate absent; posterior
nuchal plate present with bony dorsomedial crest; large gap
present between the supraoccipital and posterior nuchal
plate; one to three dorsal spines at midline of snout,
posteriormost of these spines large; one large double and
rugose lateral head spine present below the cup-like supra-
occipital crest; three small blunt lateral head spines on
operculum; rim of orbit elevated dorsolaterally and strongly
ventrally; two spines on cleithral ring; large rugose spine
anterior to ventral third of pectoral-fin base; moderate-sized
spine at ventral extent of head; small spine present
posterolateral to the pelvic-fin base; four subdorsal spines,
forming a square, the dorsal two enlarged.
Etymology.—The generic name Cylix is derived from the
Greek kylix, meaning cup or chalice, in reference to the cup-
like crest present on the head. Gender masculine.
Cylix tupareomanaia Short, Trnski, and Ng¯
atiwai, new
species
urn:lsid:zoobank.org:act:4DAADA17-DC8E-43EB-B1B8-
6495E8886C05
Common Names: M¯
aori—Tu pare o manaia, English—Manaia
Pygmy Pipehorse
Figures 1–5, Tables 1–5
Hippocampus jugumus: Kuiter, 2009: 93, figs. A, B (Poor
Knights Islands, New Zealand).
Acentronura australe: Stewart, 2015: 1053, fig. 148.1 (Bay of
Islands, New Zealand).
Idiotropiscis aotearoa: Perkins, 2017 (Whangaruru, New
Zealand; http://www.inspiredtodive.com/photo-blog/
introducing-idiotropiscis-aotearoa).
Holotype.—AIM MA122274, 31.4 mm SL, female, New
Zealand, Waiatapaua Bay, Whangaruru, 35819018.700 S,
174822008.1 00 E, depth 14 m, hand collected via SCUBA on
vertical rock wall covered in encrusting coralline algae,
bryozoans, sponges, solitary corals, turf algae, with Ecklonia
in adjacent area, C. Bedford, S. Hannam, I. Middleton, G.
Short, and T. Trnski, 11 April 2017.
Paratypes.—NMNZ P.046322, 55.5 mm SL, male, New
Zealand, Bay of Islands, east of Oturori Rock, 35814053.900 S,
174809035.1 00 E, depth 12–17 m, beam trawl, trip code
kah0907, RV Kaharoa, shallow rocky reef and soft sediments
with a mixture of Caulerpa,Ecklonia, and red and brown
algae, M. Morrison, N. Bagley, NIWA, 3 September 2009;
NMNZ P.056154, 35.5 mm SL, female, New Zealand, Cavalli
Islands, Cavalli Passage, 35800050.4 00 S, 173855026.400 E, depth
12.6–14.5 m, beam trawl, mixture of brown algae, Ecklonia,
Caulerpa,Lissonia, rhodoliths, and assorted red and brown
algae, C. and I. Middleton, NIWA, 21 April 2014.
Diagnosis.—See generic diagnosis.
Description.—Morphometric and meristic characters of the
three type specimens listed in Table 1. Trunk rings 13–14; tail
rings 35–36; anal-fin rays 4; subdorsal rings 3 (spans one
trunk ring and two tail rings); dorsal-fin rays 14; anal-fin rays
4; pectoral-fin rays 14. Body slender; head large relative to
body, angled ventrally approximately 258from the principal
body axis, the dorsal profile pyramidal in lateral aspect, rising
steeply from snout to elevated and prominent supraoccipital;
distinct cup-like crest (SC) present anterodorsally on the
supraoccipital well behind the eye, height moderate, pen-
tamerous in dorsal view, divided transversally into two
concave sections (Figs. 3, 4); cleithral and supracleithral
ridge prominent (Fig. 4); posterior margins of pentamerous
crest on supraoccipital fused and equal in height to
cleithrum; anterior nuchal plate absent; posterior nuchal
plate present with bony dorsomedial crest; large gap between
the supraoccipital and posterior nuchal plate; gill openings
small, bilateral; rim of orbit with prominent dorsolateral and
ventral ridges, fluted with rugose sculpturing; opercular ridge
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low, entire, angled dorsally toward gill opening; swelling of
gular region posteroventrally of eye, forming a transverse pair
of blunt protuberances; pectoral-fin base without distinct
ridges, one strongly elevated ventrolateral bulge (Fig. 4);
dorsal-fin origin on 12
th
trunk ring, fin base elevated;
superior trunk ridge discontinuous with superior tail ridge
below dorsal-fin base; lateral trunk ridge continuous with
inferior tail ridge; inferior tail ridge ends on anal trunk ring;
dorsum of anteriormost two trunk rings distinctly broader
than posterior trunk rings; trunk in lateral view narrowest at
1
st
and 2
nd
trunk rings where angle of head forms from body
axis, broadest at 5
th
trunk ring; principal body ridges distinct
and moderately elevated; tail rings of uniform depth over
most of length, becoming progressively shorter and smaller
near posterior tip; tail prehensile; scutella not evident.
Large spine present on dorsal midline of snout on the
ethmoid area, at confluence with the anterior ends of
supraorbital ridges, its height extended well above level of
nares; none to two smaller medial spines anterior to the large
spine on the snout, on the mesethmoid bone (principal
dorsal spine and one anterior spine in paratype NMNZ
P.056154, principal spine with anterior dorsal spines or
elevations absent in paratype NMNZ P.046322); distinct
median frontal spine at convergence of anterior edges of
the cup-like supraoccipital crest, protruding anteriorly; four
lateral head spines, one large double and rugose lateral head
spine directly below the cup-like supraoccipital crest, three
small blunt spines on operculum aligned 55–588relative to
the ventral axis of the head, the dorsal and ventral blunt
spines connecting to terminal elements of the opercular
ridge; large, conspicuous midventral conical spines on the
cleithral symphysis and the first trunk ring between the
pectoral-fin bases (Figs. 3, 5); two spines on cleithral ring,
large rugose spine anterior to ventral third of pectoral-fin
Fig. 1. Cylix tupareomanaia. (A) AIM MA122274, female, holotype shortly after death, 31.4 mm SL; Waiatapaua Bay, Whangaruru, Northland, New
Zealand (photograph ÓAuckland Museum). (B) NMNZ P.056154, female, paratype, shortly after death, 35.5 mm SL; Cavalli Islands, Northland, New
Zealand (photograph ÓIrene Middleton).
Short and Trnski—New pygmy pipehorse from New Zealand 809
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base, moderate-sized spine at ventral extent of head; small
spine present posterolaterally of the pectoral-fin base (Figs. 3,
5); superior trunk ridges with spines of small to moderate
size, enlarged blunt spines dorsally on 3
rd
,6
th
,10
th
,11
th
, and
12
th
rings, all bearing dermal flaps; lateral trunk ridges with
moderate-sized spines on each trunk ring starting at 2
nd
ring
with enlarged spines on 3
rd
,6
th
, and 10
th
rings; inferior trunk
ridges with moderate-sized spines starting at 3
rd
ring with
enlarged spines on 3
rd
,6
th
, and 10
th
rings; subdorsal spines
four, superior trunk ridge ending with two subdorsal spines,
Fig. 2. Cylix tupareomanaia. (A) AIM MA122274, female, preserved holotype, 31.4 mm SL; Waiatapaua Bay, Whangaruru, Northland, New Zealand
(photograph ÓAuckland Museum). (B) NMNZ P.056154, female, preserved paratype, 35.5 mm SL; Cavalli Islands, Northland, New Zealand
(photograph ÓAuckland Museum). (C) NMNZ P.046322, male, preserved paratype, 55.5 mm SL; east of Oturori Rock, Bay of Islands, Northland, New
Zealand (photograph Graham Short).
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the anteriormost spine large and conspicuous, superior tail
ridge commencing with two subdorsal spines, posteriormost
spine reduced, in alignment with larger subdorsal spines
above; superior tail ridge spines well developed anteriorly,
except on first and second ring, with enlarged spines on 3
rd
,
4
th
,7
th
, and 11
th
tail ridges, gradually reducing in size to 22
nd
trunk ring; lateral tail ridge spines absent; inferior tail ridge
spines well developed to 8
th
tail ring. Simple and branched
dermal appendages present on head: long simple appendages
extending anteriorly from dorsal rim of orbit, long and
branched appendages ventrodorsally of each eye.
Cylix tupareomanaia exhibits strong sexual dimorphism
with an enclosed brood pouch in male paratype NMNZ
P.046322 (Fig. 2). The brood pouch is located along the
ventral midline of the tail below the anteriormost ten tail
rings. It is enclosed by ten arcuate bony extensions (Fig. 4)
that extend ventrolaterally from the anterior ventral plate
ridges of the tail, and progressively reduce in size posteriorly.
The ten brood pouch plates are similar in appearance,
whereas the posteriormost pouch plate is diminutive in size.
Coloration.—Holotype in life (Fig. 6A), head, trunk, and tail
red; ventrolateral margin of trunk pale red to white; dorsum
of head and snout speckled with fine white dots; pentamer-
ous crest on supraoccipital red; snout spines, supraoccipital
spine, anterior continuations of supraorbital ridges, and
dorsal rim of orbit pale brown to white; white band
extending from just behind the eye, grading posteriorly into
a reticulated pattern of irregular, roundish quadrilaterals
delineated by white coloration, concentrated on operculum
and pectoral-fin base; reticulated pattern diffused laterally on
head, pale brown to white; dorsal-fin base white with
reticulated pattern, proximal third of dorsal-fin red; two
parallel rows of rounded quadrilaterals present on trunk and
tail rings, four quadrilaterals per ring; medioventral conical
spines on the cleithral symphysis and the first trunk ring
between the pectoral-fin bases pale brown to white. Fleshy
appendages, pale brown to white, present on the frontal
spine, principal snout spine, dorsum of rim of orbit,
ventrolateral of snout, 3
rd
and 6
th
superior trunk ridge
spines, and 3
rd
and 7
th
superior tail ridge spines. Other
individuals of C. tupareomanaia observed at the type location
and the Poor Knights Islands exhibited red and white
countershading coloration or background color typically
uniformly pale orange to dark red, respectively (Fig. 6B–F).
In alcohol, head and body color pale cream to light brown.
Fins hyaline.
Distribution and habitat.—Cylix tupareomanaia is thus far
known only from Taitokerau Northland, New Zealand,
including P¯
ewhairangi (Bay of Islands), Nukutaunga (Cavalli
Islands), Tawhiti Rahi and Aorangi (Poor Knights Islands),
and the coastal headlands off Whangaruru (Fig. 7). At the
type locality, C. tupareomanaia was observed in mixed habitat
on a vertical rock wall that was covered in encrusting
coralline algae, bryozoans, sponges, solitary corals, and turf
algae, and which faced a rocky outcrop exposed to the open
ocean. The rock wall was subject to moderate ocean swell as
was evident from the accumulated debris of detached and
decomposed algae on the sandy to rubble bottom of the
channel at a depth of 17–18 m. The holotype was collected
below the Ecklonia radiata kelp line; it was observed moving
about in close proximity to another individual of the same
Fig. 3. Head of Cylix tupareomanaia (AIM MA122274, female,
preserved holotype, 31.4 mm SL) in lateral (A), dorsal (B), and ventral
(C) view highlighting positions of diagnostic characters, including the
cup-like supraoccipital crest present on the supraoccipital, and large
medioventral conical spines on the cleithral symphysis and the first
trunk ring between the pectoral-fin bases. Abbreviations: CL, cleithral
spines; CSS, medioventral conical spine on the cleithral symphysis;
DHS, double head spine; FS, frontal spine; LHS, lateral head spine;
MVFTRS, medioventral first trunk ring spine between the pectoral-fin
bases; ORE, orbital rim extension; PLS, posterolateral spine on pectoral-
fin base; PNPC, posterior nuchal plate crest; SC, supraoccipital crest;
SnS, snout spines.
Short and Trnski—New pygmy pipehorse from New Zealand 811
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species, which appeared to represent a male–female pair.
Additional reef fishes observed on and at the base of the rock
wall included: Gobiopsis atrata (Gobiidae); Forsterygion flavo-
nigrum,F. maryannae, and Notoclinops segmentatus (Tripter-
ygiidae); Aplodactylus arctidens (Aplodactylidae); Coris
sandeyeri and Notolabrus tetricus (Labridae); Chromis dispilus
(Pomacentridae); and Chrysophrys auratus (Sparidae).
Species composition and biodiversity around the North
Island of New Zealand indicate subdivision of inshore marine
reef ichthyofauna into three regional biogeographic groups
or eco-regions, including: western North Island coast; north-
eastern North Island coast and offshore islands; and
Manawat¯
awhi Three Kings Islands (Brook, 2002). Cylix
tupareomanaia may be restricted to the warm temperate
waters off the north-eastern coast of the North Island and its
offshore islands where it has been collected and observed.
The coastal headlands and offshore islands are strongly
influenced by oceanic water masses and reflect higher overall
species diversity (Brook, 2002); therefore, the new taxon may
not occur in the other cooler regional biogeographic areas. Its
occurrence further north and south, however, may be
confirmed by comprehensive sampling for this cryptic
species. The new species is likely endemic to temperate
New Zealand; no observations as of yet have been recorded at
Rangit¯
ahua the Kermadec Islands, where more than 90% of
the coastal ichthyofauna are of subtropical and tropical Indo-
Pacific origin, in temperate or tropical Australia, including its
offshore subtropical territories of Lord Howe and Norfolk
Islands, nor in New Caledonia (Allen et al., 1976; Francis,
1993; Francis and Randall, 1993; Johnson, 1999; Hutchins,
2001; Gomon, 2007; Fricke et al., 2011; Larson et al., 2013;
Francis and Duffy, 2015; Trnski et al., 2015; Clark et al.,
2017).
Fig. 4. lCT scan of Cylix tupareomanaia, NMNZ P.046322, male, paratype, 55.5 mm SL. (A, B) Anterolateral view of the head highlighting the
bifurcated and cup-like crest present on the supraoccipital, continuous cleithral ring, and the strongly elevated ventrolateral bulge of the pectoral-fin
base. (C) Anterodorsal aspect of the neurocranium highlighting the bifurcated and cup-like pentamerous bony crest present on the supraoccipital.
Abbreviations: FS, frontal spine; PFB, pectoral-fin base; SC, supraoccipital crest; SCL, supracleithrum.
812 Ichthyology & Herpetology 109, No. 3, 2021
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Cryptic fish assemblages on temperate rocky reefs in New
Zealand are dominated by the highly diverse triplefins
(Tripterygiidae) and clingfishes (Gobiesocidae) and exhibit a
high degree of endemism (Paulin and Roberts, 1993; Hickey et
al., 2009; Roberts et al., 2015; Conway et al., 2017, 2018).
Other diminutive cryptic reef species recorded include the
blennies (Blenniidae), kelpfishes (Clinidae), gobies (Gobiidae),
and pipefishes and seahorses (Syngnathidae). The recent
discovery of the highly cryptic C. tupareomanaia is therefore
a noteworthy addition to reef fish diversity in New Zealand
based on the fact it hadn’t been collected or observed in
numerous biodiversity surveys of temperate marine ichthyo-
fauna previously conducted in a wide range of habitats. These
surveys were carried out in estuaries, shallow coastal bays,
seagrass meadows, mangrove forests, open mainland coasts,
peninsulas, nearshore islands, and passages within the Bay of
Islands (Nicholson, 1979; Nicholson and Roberts, 1980;
Francis et al., 2005, 2011; Kelly, 2007; Jones et al., 2009;
Gordon et al., 2010), throughout northern New Zealand
(Willan et al., 1979; Brook, 2002; Morrison et al., 2002, 2014;
Francis et al., 2005; Leathwick et al., 2006; Edgar et al., 2013),
along the coast of the East Cape Region (Roberts and Stewart,
2006), at various inshore and offshore islands, including
Manawat¯
awhi Three Kings Islands (Hardy et al., 1987), Cavalli
Islands (Nicholson, 1979), the Poor Knights Islands (Russell,
1970; Kelly, 2007), Aotea Great Barrier Island (Roberts et al.,
1986; Sivaguru and Grace, 2004), the Mokohinau Islands
(Housley, 1980), Hen Island (Willis, 1995), Cuvier Islands
(Housley et al., 1981), Ahuahu Grea Mercury Island (Grace,
1976), Aldermen Islands (Grace, 1973), the Chatham Islands
(Roberts, 1991), in the South Island (Francis, 1979; Leathwick
et al., 2006; Gordon et al., 2010; Francis et al., 2011; Morrison
et al., 2014), and the sub-Antarctic Auckland Islands (Kings-
ford et al., 1989).
Cylix tupareomanaia is currently known from very few
collected specimens and observations, which suggests that
this species occurs in low abundance throughout its range, is
hard to find due to its diminutive size and excellent crypsis,
only occasionally occurs within SCUBA depths, or is simply
rare in the regions where surveys have been conducted.
These inferences are supported by the fact that representa-
Fig. 5. lCT scan of the ventral aspect of first trunk ring of Cylix
tupareomanaia, NMNZ P.046322, male, paratype, 55.5 mm SL, in
ventral aspect highlighting positions of large medioventral conical
spines on the cleithral symphysis and the first trunk ring between the
pectoral-fin bases. Abbreviations: CL, cleithral spines; CSS, medioventral
conical spine on the cleithral symphysis; MVFTRS, medioventral first
trunk ring spine between the pectoral-fin bases; PLS, posterolateral
spine on pectoral-fin base.
Table 1. Counts and proportional measurements of type specimens of Cylix tupareomanaia. Ratios are separated by a colon and are listed as
percent of SL, HL, or SnL. Abbreviations: CH (coronet height), DL (dorsal-fin length), HD (head depth), HL (head length), OD (orbital height), PL
(pectoral-fin length), PO (post-orbital length), SL (standard length), SnD (snout depth), SnL (snout length), TaL (tail length), TD4 (trunk depth
between the 4
th
and 5
th
trunk rings), TD9 (trunk depth between the 9
th
and 10
th
trunk rings), TrL (trunk length).
Cylix tupareomanaia Cylix tupareomanaia Cylix tupareomanaia
Voucher number AIM MA122274 NMNZ P.046322 NMNZ P.056154
Type Holotype Paratype Paratype
SL (mm) 31.4 60.6 39.9
Trunk rings 13 13 14
Tail rings 36 35 36
Subdorsal trunk rings 3 3 3
Dorsal-fin rays 14 14 14
Anal-fin rays 4 4 4
Pectoral-fin rays 14 14 14
HL:SL 19.6 14.9 15.6
TrL:SL 28.4 26.6 30.8
DL:SL 5.3 5.8 5.76
PL:SL 3.12 3.0 3.1
TaL:SL 52.2 58.6 53.6
TD4:SL 7.7 8.9 7.30
TD9:SL 6.8 6.9 6.9
SnD:SnL 52.5 55.7 40.2
HD:HL 59.3 67.7 60.8
SnL:HL 32.6 42.1 39.2
OD:HL 16.3 15.7 19.4
PO:HL 42.7 40.0 43.5
CH:HL 48.2 52.6 52.2
HL:TrL 69.2 55.8 50.6
Short and Trnski—New pygmy pipehorse from New Zealand 813
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Table 2. Morphological counts and proportional measurements for type specimens of Cylix tupareomanaia and members of Acentronura and Idiotropiscis. Measurements are in percent SL and HL.
Abbreviations: CH (coronet height), H (holotype), HD (head depth), HL (head length), NT (non-type), P (paratype), PO (post-orbital length), SL (standard length), SnD (snout depth), SnL (snout
length), TaL (tail length), TD4 (trunk depth between the 4
th
and 5
th
trunk rings), TD9 (trunk depth between the 9
th
and 10
th
trunk rings), TrL (trunk length). Lines present, from top to bottom,
numbers for standard length (SL), counts for trunk rings, tail rings, dorsal and pectoral fins, and proportions.
Species C. tupareomanaia, new species I. australe I. larsonae I. lumnitzeri A. breviperula A. gracilissima A. tentaculata
Voucher AIM
MA122274
NMNZ
P.046322
NMNZ
P.056154
WAM
P.33543001
SAMA
F2657
NTM
S.10805-001
GCRL
21518
CAS
HH-0423
AMS
I.45395-001
CAS
247135
CAS-SU
6681
CAS
247139
M
CAS
247139
F
Type H P P NT NT H P NT NT NT NT NT NT
SL (mm) 31.4 60.6 39.9 54.7 50.9 33.5 33.0 69.9 54.0 40.1 70.4 50.8 53.9
Trunk rings 13 13 14 13 13 11 11 11 11 12 13 12 12
Tail rings 36 35 36 37 37 39 40 45 43 42 45 44 43
Subdorsal
trunk rings
3334 44 444 4 444
Dorsal-fin
rays
14 14 14 16 16 17 17 15 15 15 16 16 16
Pectoral-fin
rays
14 14 14 15 16 12 13 13 13 14 15 14 14
HL:SL 16.5 14.7 15.6 15.1 15.1 15.3 14.4 14.2 15.6 16.7 14.3 15.1 16.4
TrL:SL 28.3 26.6 30.8 24.3 26.3 24.8 26.2 18.6 20.1 27.9 25.1 22.6 27.7
TaL:SL 54.2 58.6 53.6 60.5 61.7 62.7 52.5 67.2 63.5 55.4 60.5 62.3 60.0
TD4:SL 7.7 8.9 7.3 7.7 6.4 12.5 12.2 5.7 6.7 8.1 6.8 6.7 8.7
TD9:SL 6.8 6.9 6.9 9.6 9.3 14.6 10.9 6.9 7.8 7.1 6.1 6.6 8.6
SnD:SnL 52.5 55.7 40.2 46.2 37.6 60.4 75.4 54.7 47.7 41.9 32.0 43.2 53.7
HD:HL 59.3 67.7 60.8 59.0 50.9 72.0 73.0 60.4 56.5 46.1 43.4 42.2 45.6
SnL:HL 32.6 42.1 39.2 31.9 38.3 29.0 24.9 33.1 31.2 36.7 35.9 35.6 33.5
PO:HL 42.7 40.6 43.5 48.0 47.4 46.1 51.3 52.3 41.6 46.1 45.2 42.8 45.3
CH:HL 48.2 52.6 52.2 34.5 43.9 66.1 67.7 39.4 44.1 36.1 38.3 29.7 32.3
HL:TrL 63.1 55.8 50.7 62.2 57.6 61.9 54.9 76.6 74.2 59.8 57.2 66.7 59.2
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tives of Idiotropiscis in southern Australia are uncommon in
its respective distributions; only two records currently exist
for I. larsonae, and seven for I. australe (OZCAM museum
record search; https://ozcam.ala.org.au/occurrences/search?
taxa¼Idiotropiscis#tab_recordsView).
Etymology.—The species epithet tupareomanaia is a neologism
gifted by kaum¯
atua (tribal elders) of Ng¯
atiwai and references
Home Point adjacent to the type locality, referred to by
Ng¯
atiwai as Tu Pare o Huia, meaning ‘‘the plume of the huia’’;
the huia was a bird that became extinct in the early 20
th
century. Tu Pare o Manaia translates as ‘‘the garland of the
Manaia.’’ The pare, or garland, references the pentamerous
head crest of the new species, and Manaia is the M¯
aori name
for a seahorse, and is also an ancestor that appears as a stylized
figure used in M¯
aori carvings representing a guardian.
Remarks.—Apparent ontogenetic differences in morphologi-
cal features of the head were observed between the
specimens of C. tupareomanaia examined herein (Fig. 2).
The smallest female (Fig. 2A, AIM MA122274, 31.4 mm SL,
holotype) exhibits a cup-like crest on the supraoccipital that
is highly elevated anteriorly, distinct median supraoccipital
spine at convergence of anterior edges of the coronet,
protruding well anteriorly; three dorsal spines at midline of
snout on mesethmoid bones, exceptionally large principal
dorsal spine at confluence with anterior continuations of
supraorbital ridges, the two anterior dorsal spines small, and
the lateral head spine directly ventral of the cup-like
supraoccipital crest consists of distinct but small paired
spines. The morphological features in the larger female
paratype (Fig. 2B, NMNZ P.056154, 35.5 mm SL) are less
pronounced: the median supraoccipital spine presents as a
distinct but blunt spine protruding anteriorly, the snout
Table 3. Comparison of meristic and morphometric characters between Cylix tupareomanaia and members of Acentronura and Idiotropiscis.
Abbreviations: HT (Holotype), NT (Non-type), PT (Paratype).
Species C. tupareomanaia, new species I. australe
Type HT PT PT NT NT
Voucher number AIM
MA122274
NMNZ
P.046322
NMNZ
P.056154
WAM
P.33543001
SAMA
F2657
Trunk rings 13 13 14 13 13
Tail rings 36 35 36 37 37
Subdorsal trunk rings 3 3 3 4 4
Dorsal-fin rays 14 14 14 16 16
Pectoral-fin rays 14 14 14 15 16
Head angled ventrad 258Present Present Present Present Present
Superior and inferior trunk and tail ridges Discontinuous Discontinuous Discontinuous Discontinuous Discontinuous
Cup-like crest on supraoccipital Present Present Present Absent Absent
Dorsomedial bony crest on supraoccipital Absent Absent Absent Present Present
Dorsomedial bony crest on first trunk ring Present Present Present Present Present
Bilateral protuberances present lateroposteriorly on
posttemporal
Absent Absent Absent Present Present
Large rugose lateral head spine Present Present Present Absent Absent
3 small lateral head spines Present Present Present Absent Absent
Frontal spine Present Present Present Absent Absent
Snout spines Present Present Present Present Present
Midventral conical spines on cleithral symphysis
and first trunk ring
Present Present Present Absent Absent
Cleithral ring confluent Present Present Present Absent Absent
Cleithral ring spine on pectoral-fin base Present Present Present Absent Absent
Cleithral ring spine ventrally of first trunk ring Present Present Present Absent Absent
Small spine ventroposteriorly of pectoral-fin base Present Present Present Absent Absent
Subdorsal spines Present Present Present Absent Absent
Orbital rim extension Present Present Present Present Present
Swelling of gular region ventroposterior to eye Present Present Present Present Present
Superior trunk ridge spines 3, 6, 10 Absent Absent
Superior tail ridge spines 4, 7, 11 Absent Absent
Anal-fin Present Present Present Present Present
Table 4. Summary of the number of trunk rings spanning the brood pouch in male specimens of C. tupareomanaia and members of Acentronura
and Idiotropiscis.
Species C. tupareomanaia A. breviperula A. gracilissima A. tentaculata I. australe I. larsonae I. lumnitzeri
Voucher NMNZ
P.046322
BMNH
1890.1.14.51
CAS-SU
6681
BMNH
1869.6.21.7
WAM
P. 33543.001
NTM
S.10805-001
AMS
I.38660-001
Type Paratype Holotype Non-type Holotype Non-type Holotype Holotype
Trunk rings 8 7 14 12 10 8 11
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possesses two dorsal spines, the principal spine and one small
anterior spine, and the lateral head spine directly ventral of
the bony pentamerous bony crest consists of small but
merged paired spines. In contrast, the adult male paratype
(Fig. 2C, NMNZ P.046322, 55.5 mm SL) exhibits a cup-like
supraoccipital crest that is low, less elevated, angled some-
what posteriorly, and cup-like in dorsal view, the median
frontal spine a minute but discernible ridge, wedge-like in
appearance, one large principal spine at midline of snout,
and the lateral head spine directly ventral of the pentamer-
ous bony crest is merged as one rugose spine.
Acentronura Kaup, 1853
Diagnosis.—A genus of the Syngnathidae that shares numer-
ous morphological synapomorphies with Cylix and Idio-
tropiscis, including head angled ventrally approximately 258
from the principal body axis, enclosed brood pouch, brood
Table 3. Extended.
I. larsonae I. lumnitzeri A. breviperula A. gracilissima A. tentaculata
HT PT NT NT NT NT NT NT
NTM
S.10805-001
GCRL
21518
CAS
HH-0423
AMS
I.45395-001
CAS
247135
CAS-SU
6681
CAS
247139
CAS
247139
11 11 11 11 12 13 12 12
39 40 45 43 42 45 44 43
4444 4444
17 17 15 15 15 16 16 16
12 13 13 13 14 15 14 14
Present Present Present Present Present Present Present Present
Discontinuous Discontinuous Discontinuous Discontinuous Continuous Continuous Continuous Continuous
Absent Absent Absent Absent Absent Absent Absent Absent
Present Present Present Present Present Present Present Present
Present Present Present Present Present Present Present Present
Present Present Present Present Present Present Present Present
Absent Absent Absent Absent Absent Absent Absent Absent
Absent Absent Absent Absent Absent Absent Absent Absent
Absent Absent Absent Absent Absent Absent Absent Absent
Absent Absent Absent Absent Absent Absent Absent Absent
Absent Absent Absent Absent Absent Absent Absent Absent
Absent Absent Absent Absent Absent Absent Absent Absent
Absent Absent Present Present Absent Absent Absent Absent
Absent Absent Absent Absent Absent Absent Absent Absent
Absent Absent Present Present Absent Absent Absent Absent
Absent Absent Present Present Absent Absent Absent Absent
Absent Absent Present Present Absent Absent Absent Absent
Present Present Present Present Absent Absent Absent Absent
Absent Absent 6, 10 Absent Absent Absent Absent
Absent Absent Absent Absent Absent Absent Absent Absent
Present Present Present Present Present Present Present Present
Table 5. Morphological character comparisons between Hippocampus and the pygmy pipehorse genera Cylix and Idiotropiscis.
Species Hippocampus spp. C. tupareomanaia I. australe I. larsonae I. lumnitzeri
Coronet Present Present Absent Absent Absent
Subdorsal rings 4 Present Present Present Present Present
Parietal spine Present Present Absent Absent Absent
Lateral head spines ventral of coronet Present Present Absent Absent Absent
Snout spine Present Present Present Present Absent
Cleithral ring spines Present Present Absent Absent Present
Subdorsal spines Present Present Absent Absent Present
Enlarged superior trunk spines Present Present Absent Absent Absent
Enlarged superior tail spines Present Present Absent Absent Absent
Enlarged spines on lateral and
inferior trunk spines
Present Present Absent Absent Absent
Eye spines Present Absent Absent Absent Absent
Elevated rims of orbit Absent Present Absent Absent Present
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pouch plates, prehensile tail, and absence of caudal fin.
However, Acentronura differs from its co-familials by the
following combination of characters: supraoccipital low, not
elevated, with distinct dorsomedial crest, followed by an
elevated bony dimple; bilateral bony lobed protuberances on
the posterolateral margins of the post-temporal bones;
anterior nuchal plate absent; posterior nuchal plate present
anterior to cleithrum with dorsomedial crest-like ridge along
its dorsum, large gap present between the supraoccipital and
posterior nuchal plate; small blunt spine midway between
orbit and lobed protuberance on operculum; cleithral ring
distinct, discontinuous mid-dorsally; single gill slit between
supraoccipital and cleithral ridge; rim of orbit projecting
dorsolaterally and slightly ventrolaterally; spines absent on
principal trunk and tail ridges.
Fig. 6. Cylix tupareomanaia in situ. (A) AIM MA122274, female, holotype, Waiatapaua Bay, Whangaruru, Northland, New Zealand, 12 m depth
(photograph ÓShane Housham). (B) Waiatapaua Bay, Whangaruru, Northland, New Zealand, 12 m depth (photograph ÓShane Housham). (C)
Waiatapaua Bay, Whangaruru, Northland, New Zealand, 12 m depth (photograph ÓRichard Smith). (D) Waiatapaua Bay, Whangaruru, Northland,
New Zealand, 12 m depth (photograph ÓIrene Middleton). (E) Waiatapaua Bay, Whangaruru, Northland, New Zealand, 12 m depth (photograph Ó
Irene Middleton). (F) Poor Knights Islands, Northland, New Zealand, at 10 m depth (photograph ÓKent Erickson).
Short and Trnski—New pygmy pipehorse from New Zealand 817
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Type species.—Hippocampus gracilissimus Temminck and
Schlegel, 1850.
Acentronura breviperula Fraser-Brunner and Whitley,
1949
Figures 8A, 9A, 10A, 11A; Tables 2–4
Acentronura breviperula Fraser-Brunner and Whitley, 1949:
148, fig. 1 (Mabuiag, Torres Strait, Queensland).
Diagnosis.—See generic diagnosis. Acentronura breviperula
differs from its congeners by the following combination of
characters: supraoccipital consisting of a distinct bony
bipartite dorsomedial crest, segments arcuate in lateral view,
followed by an elevated and long bony dimple; posterior
margin of second segment of dorsomedial crest merging
within the bony dimple, which is approximately one-third
longer than the dorsomedial crest and tapered posteriorly.
Description.—Morphometric and meristic characters listed in
Tables 2–4. Superior trunk and tail ridges discontinuous
below dorsal-fin base, lateral tail ridge present, inferior trunk
ridge ends at anal ring, lateral trunk ridge confluent with
inferior tail ridge. Head angled ventrally 258from the
principal body axis, the anterodorsal profile slightly conoid
in lateral aspect; supraoccipital low, not elevated, with a
distinct bony bipartite dorsomedial crest, segments arcuate in
lateral view, followed by an elevated and long bony dimple;
posterior margin of second segment of dorsomedial crest
merging within the bony dimple, which is approximately
one-third longer than the dorsomedial crest and tapered
posteriorly; bilateral bony lobed protuberances on the
posterolateral margins of the post-temporal bones (Fig.
10A); anterior nuchal plate absent (Fig. 9A); posterior nuchal
plate present anterior to cleithrum with dorsomedial crest-
like ridge along its dorsum, large gap between the supraoc-
cipital and posterior nuchal plate; small blunt spine midway
between orbit and lobed protuberance on operculum;
cleithral ring distinct, discontinuous mid-dorsally; gill slit
between supraoccipital and cleithral ridge; rim of orbit
projecting dorsolaterally and slightly ventrolaterally; snout
spine absent; interorbital narrow, depressed; opercular ridge
distinct, entire, angled dorsally toward gill opening; swelling
of gular region ventroposterior to orbit absent; pectoral-fin
base without distinct ridges, low, strongly elevated ventro-
lateral bulge supporting the pectoral-fin absent; trunk
deepest anteriorly, principal body ridges distinct; principal
body ridge spines absent; caudal fin absent. Acentronura
breviperula exhibits strong sexual dimorphism associated
with the presence of a brood pouch. The brood pouch is
formed along the ventral midline of the tail and is present
below the anteriormost eight tail rings (Fig. 11A).
Distribution.—Acentronura breviperula is known from the
tropical Indo-West Pacific, from East Africa, Madagascar, the
Red Sea, Persian Gulf, Papua New Guinea, Torres Strait,
northern Great Barrier Reef, Australia, New Caledonia, and
Taiwan (Fraser-Brunner and Whitley, 1949; Dawson, 1985;
Kuiter, 2000, 2003; Fricke and Kulbicki, 2006; Chen, 2017;
Fricke et al., 2019).
Material examined.—Acentronura breviperula: CAS 247135,
female, 40.1 mm SL, channel between Iles aux Canards and
Noumea,AnseVateBay,NewCaledonia,22818023.800 S,
166826007.9 00 E, depth 23 m, H. Hamilton and G. Short, 18
October 2008.
Acentronura gracilissima Temminck and Schlegel, 1850
Figures 8B, 9B; Tables 2–4
Hippocampus gracilissimus Temminck and Schlegel, 1850: 274,
pl. 120, fig. 6 (Japan).
Atelurus germani Dum´
eril, 1870: 584 (Cochin, China).
Diagnosis.—See generic diagnosis. Acentronura gracilissima
differs from its congeners by the following combination of
characters: supraoccipital consisting of a distinct dorsomedial
crest, non-segmented, ridge-like in lateral view, followed by
an elevated and bony dimple; bony dimple approximately
one-third in length of dorsomedial crest.
Fig. 7. Distribution of Cylix tupareo-
manaia in Taitokerau Northland,
North Island, New Zealand. Square
¼type locality of holotype AIM
MA122274 at Waiatapaua Bay,
Whangaruru. Star ¼locality of para-
type NMNZ P.056154, Cavalli Islands.
Circle ¼locality of paratype NMNZ
P.046322, east of Oturori Rock, Bay
of Islands. Arrow ¼specimen photo-
graphed at Poor Knights Islands.
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Fig. 8. Lateral view of preserved specimens of Acentronura spp. redescribed in this study. (A) A. breviperula, CAS 247135, female, 40.1 mm SL. (B)
A. gracilissima, CAS-SU 6681, male, 70.4 mm SL. (C) A. tentaculata, CAS 247139, male, 50.8 mm SL. (D) A. tentaculata, CAS 247139, female, 53.9
mm SL.
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Description.—Morphometric and meristic characters listed in
Tables 2–4. Superior trunk and tail ridges discontinuous
below dorsal-fin base, lateral tail ridge present, inferior trunk
ridge ends at anal ring, lateral trunk ridge confluent with
inferior tail ridge. Head angled ventrally 258from the
principal body axis, the anterodorsal profile slightly conoid
in lateral aspect; supraoccipital, low, not elevated, with a
distinct dorsomedial crest, non-segmented, ridge-like in
lateral view, followed by an elevated and bony dimple; bony
dimple approximately one-third in length of dorsomedial
Fig. 9. Lateral view of lCT scanned skeletons of preserved specimens of Acentronura spp. redescribed in this study. (A) A. breviperula, CAS 247135,
female, 40.1 mm SL. (B) A. gracilissima, CAS-SU 6681, male, 70.4 mm SL. (C) A. tentaculata, CAS 247139, male, 50.8 mm SL. (D) A. tentaculata, CAS
247139, female, 53.9 mm SL.
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Fig. 10. Dorsal view of lCT scanned neurocranium of (A) Acentronura breviperula, CAS 247135, female; (B) Acentronura gracilissima, CAS 247139,
male; (C) Acentronura tentaculata, CAS-SU 6681, male. Abbreviations: BLP, bilateral lobed protuberances; SC, supraoccipital crest.
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Fig. 11. X-ray radiography of (A) A. breviperula, BMNH 1890.1.14.51, male, holotype; (B) A. tentaculata, BMNH 1869.6.21.7, male, holotype; (C) I.
larsonae, NTM S.10805-001, male, holotype, 55.5 mm SL (Photograph credits for A. breviperula and A. tentaculata to Oliver Crimmen and Ralf Britz,
ÓThe Trustees of the Natural History Museum, London; I. larsonae to Michael Hammer, ÓMuseum and Art Gallery of the Northern Territory).
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crest; bilateral bony lobed protuberances on the posterolat-
eral margins of the post-temporal bones (Fig. 10B); anterior
nuchal plate absent (Fig. 9B); posterior nuchal plate present
anterior to cleithrum with dorsomedial crest-like ridge along
its dorsum, large gap present between the supraoccipital and
posterior nuchal plate; small blunt spine midway between
orbit and lobed protuberance on operculum; cleithral ring
distinct, discontinuous mid-dorsally; single gill slit between
supraoccipital and cleithral ridge; rim of orbit projecting
dorsolaterally and slightly ventrolaterally; snout spine ab-
sent; interorbital narrow, depressed; opercular ridge distinct,
entire, angled dorsally toward gill opening; swelling of gular
region ventroposterior to orbit absent; pectoral-fin base
without distinct ridges, low, strongly elevated ventrolateral
bulge supporting the pectoral-fin absent; trunk deepest
anteriorly, principal body ridges distinct; principal body
ridge spines absent; caudal fin absent. Acentronura gracilissima
exhibits strong sexual dimorphism associated with the
presence of a male brood pouch. The brood pouch is formed
along the ventral midline of the tail and is present below the
anteriormost 14 tail rings (Fig. 8B). lCT scanning of the
specimen (Fig. 9B) revealed the brood pouch is enclosed by
14 arcuate bony extensions, which extend ventrolaterally
from the anterior ventral plate ridges of the tail and are
reduced in size posteriorly. The anteriormost pouch plate is
broad and paddle-shaped at ventrocaudal margin and curved
posterolaterally relative to other pouch plates. The second
pouch plate appears to consist of double arcuate bony
extensions merged as one bony extension. The posteriormost
pouch plate is diminutive in size.
Distribution.—Acentronura gracilissima is known from the
tropical western Pacific from New Caledonia, Japan, Indo-
china, and the South China Sea (Dawson, 1985; Rivaton and
Richer de Forges, 1990; Randall and Lim, 2000; Nakae et al.,
2018; Araki et al., 2019).
Material examined.—Acentronura gracilissima: CAS-SU 6681,
70.4 mm SL, Honshu Island, Sagami Sea, Japan,
35809034.0 00 N, 139829037.900 E, date collected 1900.
Acentronura tentaculata G¨
unther, 1870
Figures 8C–D, 9C–D, 11B; Tables 2–4
Acentronura tentaculata G¨
unther, 1870: 516 (Suez, Egypt).
Syngnathoides algensis Fourmanoir, 1954: 210 (Dzaoudzi,
Mayotte Island, Commore Island).
Acentronura mossambica Smith, 1963: 522, pl. 76, figs. f, g
(Inhaca and Inhambane Estuary, Mozambique).
Diagnosis.—See generic diagnosis. Acentronura tentaculata
differs from its congeners by the following combination of
characters: supraoccipital consists of distinct bony bipartite
dorsomedial crest, segments arcuate in lateral view, followed
by an elevated and bony dimple; posterior margin of second
segment of dorsomedial crest located at opening of, but not
merged within, the bony dimple, which is approximately
equal in length to the dorsomedial crest and tapered
posteriorly.
Description.—Morphometric and meristic characters listed in
Tables 2–4. Superior trunk and tail ridges discontinuous
below dorsal-fin base, lateral tail ridge present, inferior trunk
ridge ends at anal ring, lateral trunk ridge confluent with
inferior tail ridge. Head angled ventrally 258from the
principal body axis, the anterodorsal profile slightly conoid
in lateral aspect; supraoccipital, low, not elevated, with a
distinct bony bipartite dorsomedial crest, segments arcuate in
lateral view, followed by an elevated and bony dimple;
posterior margin of second segment of dorsomedial crest
located at opening of, but not merged within, the bony
dimple, which is approximately equal in length to the
dorsomedial crest and tapered posteriorly; bilateral bony
lobed protuberances on the posterolateral margins of the
post-temporal bones (Fig. 10C); anterior nuchal plate absent
(Fig. 9C–D); posterior nuchal plate present anterior to
cleithrum with dorsomedial crest-like ridge along its dorsum,
large gap present between the supraoccipital and posterior
nuchal plate; small blunt spine midway between orbit and
lobed protuberance on operculum; cleithral ring distinct,
discontinuous mid-dorsally; single gill slit between supraoc-
cipital and cleithral ridge; rim of orbit projecting dorsolater-
ally and slightly ventrolaterally; snout spine absent;
interorbital narrow, depressed; opercular ridge distinct,
entire, angled dorsally toward gill opening; swelling of gular
region ventroposterior to orbit absent; pectoral-fin base
without distinct ridges, low, strongly elevated ventrolateral
bulge supporting the pectoral-fin absent; trunk deepest
anteriorly, principal body ridges distinct; principal body
ridge spines absent; caudal fin absent. Acentronura tentaculata
exhibits strong sexual dimorphism associated with the
presence of a male brood pouch. The brood pouch is formed
along the ventral midline of the tail and is present below the
anteriormost 12 tail rings (Fig. 8C–D). lCT scanning of the
material (Fig. 9C–D) revealed the brood pouch is enclosed by
12 arcuate bony extensions, which extend ventrolaterally
from the anterior ventral plate ridges of the tail and are
reduced in size posteriorly. The anteriormost pouch plate is
broad and paddle-shaped at ventrocaudal margin and curved
posterolaterally relative to other pouch plates. The posteri-
ormost pouch plate is diminutive in size.
Distribution.—Acentronura tentaculata is known from the
tropical Indo-West Pacific, from East Africa, Madagascar, the
Red Sea, Arabian and Persian gulfs to Torres Strait, Philip-
pines, the northern Great Barrier Reef, Australia, and New
Caledonia (Dawson, 1985; Rivaton and Richer de Forges,
1990; McKenna, 2003; Fricke and Kulbicki, 2006; Al-Jufaili et
al., 2010; Grandcourt, 2012; Nakae et al., 2018).
Material examined.—Acentronura tentaculata: CAS 247139,
male, 50.8 mm SL, female, 53.9 mm SL, Tamaraw Beach,
South Puerto Galera, Philippines, 13830003.800 N,
120853040.2 00 E, Hearst Philippine Biodiversity Expedition,
station PG-011, B. Moore and H. Hamilton, 3 June 2011.
Idiotropiscis Whitley, 1947
Diagnosis.—A genus of the Syngnathidae that shares numer-
ous morphological synapomorphies with Acentronura and
Cylix, including head angled ventrally approximately 258
from the principal body axis, enclosed brood pouch, brood
pouch plates, prehensile tail, and absence of caudal fin.
However, Idiotropiscis differs from its co-familials by the
following combination of characters: supraoccipital is ele-
vated with a distinct and raised bony dorsomedial crest;
posterolateral margins of post-temporal bones expanded
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moderately anteriorly taking the form of protuberances;
anterior nuchal plate absent; posterior nuchal plate present
anterior to cleithrum with dorsomedial crest-like ridge along
its dorsum, large gap present between the supraoccipital and
posterior nuchal plate; small blunt spine midway between
orbit and lobed protuberance on operculum; cleithral ring
distinct, discontinuous mid-dorsally; bilateral gill slits situat-
ed laterally on head between supraoccipital and cleithrum.
Idiotropiscis australe (Waite and Hale, 1921)
Figures 12, 13; Tables 2–5
Acentronura australe Waite and Hale, 1921: 317–318, fig. 53
(St. Vincent Gulf, South Australia).
Diagnosis.—See generic diagnosis. Idiotropiscis australe differs
from its congeners by the following combination of
characters: distinct raised bony multipartite dorsomedial
crest on the supraoccipital, the four segments arcuate in
lateral view, spanning the complete frontal from anterior
margins of the interorbital to the supraoccipital, protruding
anteriorly; posterolateral margins of post-temporal bones
expanded moderately anteriorly taking the form of curved
protuberances; small dorsal snout spine present medially on
mesethmoid bone.
Description.—General body shape as in Figure 13. Morpho-
metric and meristic characters listed in Tables 2–4. Superior
trunk and tail ridges discontinuous below dorsal-fin base,
lateral tail ridge present, inferior trunk ridge ends at anal
ring, lateral trunk ridge confluent with inferior tail ridge.
Head angled ventrally 258from the principal body axis, the
anterodorsal profile slightly conoid in lateral aspect, rising
from short snout to an elevated supraoccipital; distinct
raised bony multipartite dorsomedial crest on the supraoc-
cipital, the four segments arcuate in lateral view, spanning
the complete frontal from anterior margins of the interor-
bital to the supraoccipital, protruding anteriorly; postero-
lateral margins of post-temporal bones expanded
moderately anteriorly taking the form of curved protuber-
ances (Fig. 14A); anterior nuchal plate absent (Fig. 13A);
posterior nuchal plate present anterior to cleithrum with
dorsomedial crest-like ridge along its dorsum, large gap
present between the supraoccipital and posterior nuchal
plate; small blunt spine midway between orbit and lobed
protuberance on operculum; cleithral ring distinct, discon-
tinuous mid-dorsally; gill openings as small bilateral slits
directly situated laterally on head between supraoccipital
and cleithrum; small dorsal snout spine present medially on
mesethmoid bone; interorbital narrow, depressed; opercular
ridge low, entire, angled dorsally toward gill opening; slight
swelling of gular region ventroposterior to eye; pectoral-fin
base without distinct ridges, on strongly elevated ventrolat-
eral bulge; trunk deepest anteriorly, principal body ridges
distinct; principal body ridge spines absent. Caudal fin
absent. Idiotropiscis australe exhibits strong sexual dimor-
phism associated with the presence of a male brood pouch.
The brood pouch is formed along the ventral midline of the
tail and is present below the anteriormost 11 tail rings (Fig.
12A). lCT scanning of the material (Fig. 13A) revealed the
brood pouch is enclosed by 11 arcuate bony extensions,
which extend ventrolaterally from the anterior ventral plate
ridges of the tail and are reduced in size posteriorly. The
anteriormost pouch plate is broad and paddle-shaped at
ventrocaudal margin and curved posterolaterally relative to
other pouch plates. The posteriormost pouch plate is
diminutive in size.
Distribution.—Idiotropiscis australe is endemic to the temper-
ate waters of southern and southwestern Australia, from
Fig. 12. Lateral view of preserved specimens of Idiotropiscis spp.
redescribed in this study. (A) I. australe, WAM P.33543-001, male, 54.7
mm SL. (B) I. larsonae, NTM S.10805-001, male, holotype, 33.5 SL. (C)
I. larsonae, GCRL 21518, female, paratype, 33.0 mm SL. (D) I.
lumnitzeri, CAS HH-0423, male, 69.9 mm SL. (E) I. lumnitzeri, AMS
I.45395-001, female, 54.0 mm SL.
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Fig. 13. Lateral view of lCT scanned skeletons of preserved specimens of Idiotropiscis spp. redescribed in this study. (A) I. australe, WAM P.33543-
001, male, 54.7 mm SL. (B) I. larsonae, GCRL 21518, female, paratype, 33.0 mm SL. (C) I. lumnitzeri, CAS HH-0423, male, 69.9 mm SL. (D) I.
lumnitzeri, AMS I.45395-001, female, 54.0 mm SL.
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Fig. 14. Dorsal view of lCT scanned neurocranium of the head of (A) Idiotropiscis australe, WAM P.33543-001, male; (B) Idiotropiscis larsonae,
GCRL 21518, male; (C) Idiotropiscis lumnitzeri, CAS HH-0423, male. Abbreviations: BLP, bilateral lobed protuberances; SC, supraoccipital crest.
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Cape Jarvis and the Gulf St Vincent, South Australia, and
Carnac Island, Western Australia (Dawson, 1985).
Material examined.—Idiotropiscis australe: SAMA F2657, Yan-
kalilla, South Australia, Australia, 35.5977678S, 138.0975118E,
depth 20 m; WAM P.25346-015, Australia, Western Australia,
Carnac Island, Gage Roads, dredge, J. Scott, 30 June 1975;
WAM P.33542-001, Success Bank, Cockburn Sound, Western
Australia, 32.0830978S, 115.6833448E, depth 4 m, 3 No-
vember 1997; WAM P.33543-001, male, Owen Anchorage,
Cockburn Sound, Western Australia, Australia, 32.1144598S,
115.7499898E, depth 12 m, 1 November 1997.
Idiotropiscis larsonae (Dawson, 1984)
Figures 11–13, Tables 2–5
Acentronura (Idiotropiscis)larsonae Dawson, 1984: 157–158,
figs. 1, 2 (Alpha Island, Monte Bello Islands, Western
Australia).
Diagnosis.—See generic diagnosis. Idiotropiscis larsonae differs
from its congeners by the following combination of
characters: bony dorsomedial crest on the supraoccipital,
strongly elevated, tripartite, the three large segments arcuate
in lateral view, spanning the complete frontal from anterior
margins of the interorbital to the supraoccipital, compressed
laterally, protruding anteriorly; posterolateral margins of
post-temporal bones expanded slightly anteriorly taking the
form of flat protuberances, not prominent; rim of orbit
projecting slightly dorsolaterally and ventrolaterally; small
dorsal snout spine present medially on mesethmoid bone;
body compressed laterally.
Description.—Morphometric and meristic characters listed in
Tables 2–4. Superior trunk and tail ridges discontinuous
below dorsal-fin base, lateral tail ridge present, inferior trunk
ridge ends at anal ring, lateral trunk ridge confluent with
inferior tail ridge. Head angled ventrally 258from the
principal body axis, the anterodorsal profile slightly conoid
in lateral aspect; bony dorsomedial crest on the supraoccip-
ital, strongly elevated, tripartite, the three large segments
arcuate in lateral view, spanning the complete frontal from
anterior margins of the interorbital to the supraoccipital,
compressed laterally, protruding anteriorly; posterolateral
margins of post-temporal bones expanded slightly anteriorly
taking the form of flat protuberances, not prominent and
difficult to discern (Fig. 14B); anterior nuchal plate absent;
posterior nuchal plate present anterior to cleithrum with
dorsomedial crest-like ridge along its dorsum, large gap
present between the supraoccipital and posterior nuchal
plate; small blunt spine midway between orbit and lobed
protuberance on operculum; cleithral ring distinct, discon-
tinuous mid-dorsally; bilateral gill slits situated laterally on
head between supraoccipital and cleithrum; median dorsal
snout spine present on mesethmoid bone; interorbital
narrow, depressed; rim of orbit projecting moderately
dorsolaterally and ventrally; opercular ridge low, entire,
angled dorsally toward gill opening; swelling of the gular
region ventroposteriorly of eye absent; pectoral-fin base
without distinct ridges, low, strongly elevated ventrolateral
bulge supporting the pectoral-fin base absent; trunk com-
pressed laterally, span of dorsum of pre-dorsal trunk rings less
by half of dorsum of anterior post-dorsal tail rings, span of
trunk at lateral and inferior ridges somewhat greater than
span of dorsum of trunk; subdorsal rings four (spans three
trunk rings and one tail rings vs. two trunk rings and two tail
rings in paratype); principal body ridges distinct; superior tail
ridge with slightly enlarged spines on 4
th
,12
th
, and 16
th
trunk rings. Caudal fin absent.
Idiotropiscis larsonae exhibits strong sexual dimorphism
associated with the presence of a brood pouch in the male
(Figs. 11C, 12B). The brood pouch is formed along the
ventral midline of the tail and is present below the anterior-
most nine tail rings. X-ray image of the holotype revealed the
brood pouch is enclosed by nine arcuate bony extensions
and reduced in size posteriorly. The female paratype exhibits
well-developed ossification of the skeleton, including the
strong ossification of the inferior and ventral trunk area as
detected by lCT scanning (Fig. 13B).
Distribution.—Idiotropiscis larsonae is known only from the
tropical waters of the Monte Bello Islands, north Western
Australia (Dawson, 1985).
Material examined.—Idiotropiscis larsonae: NTM S.10805-001,
adult male, holotype, Monte Bello Islands, Western Australia,
Australia,208360S, 1158370E, depth 3–9 m, 22 April 1983;
GCRL 21518, female, paratype, 33.0 mm SL, Monte Bello
Islands, Western Australia, Australia, 208360S, 1158370E,
depth 3–9 m, 22 April 1983.
Idiotropiscis lumnitzeri Kuiter, 2004
Figures 12–13, 15; Tables 2–5
Idiotropiscis lumnitzeri Kuiter, 2004: 165, fig. 1 (Henrietta
Head, La Perouse, Sydney, New South Wales, Australia);
AMS I.38660-001 (holotype).
Diagnosis.—See generic diagnosis. Idiotropiscis lumnitzeri
differs from its congeners by the following combination of
characters: strongly elevated supraoccipital; large, promi-
nent, and raised bony dorsomedial crest on the supraoccip-
ital, dome-like in lateral view, protruding anteriorly;
posterolateral margins of post-temporal bones expanded
moderately anteriorly taking the form of tapered and wing-
like protuberances; rim of orbit projecting dorsolaterally; one
large and blunt spine on cleithral at level of last pectoral-fin
ray; small spine posterolateral to pectoral fin; superior trunk
ridge with enlarged spines projecting dorsolaterally on 6
th
and 10
th
trunk rings; subdorsal spines four, last two superior
trunk ridges ending under dorsal fin with subdorsal spines,
first two superior tail ridges under dorsal fin with subdorsal
spines.
Description.—Morphometric and meristic characters listed in
Tables 2–4. Superior trunk and tail ridges discontinuous
below dorsal-fin base, lateral tail ridge present, inferior trunk
ridge ends at anal ring, lateral trunk ridge confluent with
inferior tail ridge. Head large relative to body, angled
ventrally 258from the principal body axis, the anterodorsal
profile conoid in lateral aspect; rising steeply from short
snout to a prominent supraoccipital; large, prominent, and
raised bony dorsomedial crest on the supraoccipital, dome-
like in lateral view, dorsocaudal margin irregular, protruding
anteriorly; posterolateral margins of post-temporal bones
expanded moderately anteriorly taking the form of tapered
and wing-like protuberances (Fig. 14C); cleithral ring distinct,
prominent, discontinuous mid-dorsally; anterior nuchal
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plate absent; posterior nuchal plate present anterior to
cleithrum with dorsomedial crest-like ridge along its dorsum,
large gap present between the supraoccipital and posterior
nuchal plate; cleithral ring distinct, low, discontinuous mid-
dorsally at bony dorsomedial crest; rim of orbit projecting
dorsolaterally; opercular ridge low, entire, angled dorsally
toward gill opening; swelling of gular region ventroposterior
to eye forming a transverse pair of blunt protuberances;
pectoral-fin base without distinct ridges, on strongly elevated
ventrolateral bulge (Fig. 15); trunk narrowest at 3
rd
and 4
th
ring; dorsum of 2
nd
,3
rd
, and 4
th
trunk rings noticeably
broader than 5–11
th
trunk rings; dorsal-fin base strongly
raised dorsally; subdorsal rings four (spans two and a half
trunk rings and one and a half tail rings), dorsal-fin base
starting immediately posterior to 10
th
trunk ring and ending
immediately posterior to 2
nd
tail ring; caudal fin absent. Body
spines: large spine present on cleithral ring at level of last
pectoral-fin ray (Fig. 15A); small spine present posterolateral
to the pelvic fin (Fig. 15A); superior trunk ridge with enlarged
spines projecting dorsolaterally on 6
th
and 10
th
trunk rings;
subdorsal spines four, last two superior trunk ridges ending
under dorsal fin with subdorsal spines, first two superior tail
ridges under dorsal fin with subdorsal spines (Fig. 15B); large
simple and branched dermal flaps present on anterior part of
frontal ridge, ventral to eye, 4
th
and 7
th
superior trunk ridges,
7–9
th
lateral and inferior trunk ridges, and on 5
th
,10
th
,15
th
,
19
th
, and 24
th
superior and inferior tail ridges. The female
specimen (AMS I.45395-001) exhibits well-developed ossifi-
cation of the skeleton, including the strong ossification of
the inferior and ventral trunk area as detected by lCT
scanning (Fig. 13D). Idiotropiscis lumnitzeri exhibits strong
sexual dimorphism associated with the presence of a male
brood pouch. The brood pouch is formed along the ventral
midline of the tail and is present below the anteriormost 11
tail rings (Fig. 12D). lCT scans revealed the brood pouch is
enclosed by 11 arcuate bony extensions (Figs. 13C, 15B),
which extend ventrolaterally from the anterior ventral plate
ridges of the tail, are reduced in size posteriorly, the
anteriormost extension and curved posterolaterally relative
to the other extensions.
Distribution.—Idiotropiscis lumnitzeri is endemic to the tem-
perate waters of southeastern Australia, from Cabbage Tree
Bay, Manly, to Brush Island located south of Ulladulla
Harbour, New South Wales (Kuiter, 2004).
Material examined.—Idiotropiscis lumnitzeri: AMS I.45395-001,
female, 54.0 mm SL, N of Burrewarra Point, S of Batemans
Bay, New South Wales, Australia, depth 22 m; CAS HH-0423,
male, 69.9 mm SL, SW of Bare Island, Botany Bay, NSW,
Australia, 33859033.3 00 S, 151813047.300 E, depth 18 m, 1 March
2007.
DISCUSSION
All previous descriptions of pygmy pipehorses are based only
on one or two specimens in which the internal (skeletal)
anatomy of the described species was undocumented. (e.g.,
Temminck and Schlegel, 1850; G¨
unther, 1870; Waite and
Hale, 1921; Fraser-Brunner and Whitley, 1949; Dawson,
1984; Kuiter, 2004). Our description of C. tupareomanaia is
also derived from a small number of individuals (holotype
and two paratypes), but contrary to these previous descrip-
tions, we have observed elements in the skeleton of C.
tupareomanaia using micro-computed tomography from
which we have identified key diagnostic features, especially
of the neurocranium (see diagnosis). The morphological
details would have been impossible to obtain from tradition-
al radiographs alone.
Comparative morphological studies among pygmy pipe-
horses.—Cylix tupareomanaia and members of Acentronura
and Idiotropiscis (Figs. 10–17), despite being superficially
similar in outward appearance, exhibit significant morpho-
logical differences among them in the neurocranium that aid
in their diagnoses. The main neurocranial distinctions were
observed in the morphology of the supraoccipital crests and
bilateral bony lobed protuberances on the posterolateral
margins of the post-temporal bones. The characters that
distinguish Cylix tupareomanaia from the members of Idio-
tropiscis are presented in Tables 2–5 and summarized below.
Cylix tupareomanaia is unique in Syngnathidae in possessing
a highly derived bony cup-like crest present anterodorsally
on the supraoccipital and large, conspicuous conical spines
ventrally centered on the head on the cleithral symphysis
and the first trunk ring between the pectoral-fin bases, which
we propose as apomorphies for this species. Cylix tupareoma-
naia is most similar to members of Idiotropiscis (Tables 2–4) in
meristic and morphometric characters and in the following
morphological and neurocranial characters: principal ridge
configuration; elevated supraoccipital; absence of anterior
nuchal plate presence of posterior nuchal plate, with large
gap between it and the supraoccipital; presence of a bony
dorsomedial crest on the dorsum of the posterior nuchal
plate; pronounced ventrolateral bulge supporting the pecto-
ral-fin base; and certain body spines (snout spines in I.
australe and I. larsonae; four subdorsal spines, large cleithral
ring spine at level of last pectoral-fin ray, and small spine
present posterolateral to the pelvic fin in I. lumnitzeri [Fig.
14]).
Cylix tupareomanaia differs primarily from the members of
Idiotropiscis by features of the crest morphology on the
supraoccipital bone (cup-like pentamerous crest [Figs. 1–4]
vs. the presence of a thin bony dorsomedial crest [Figs. 12–
15]) and large midventral spines on the cleithral symphysis
and the first trunk ring between the pectoral-fin bases (Figs.
4–6; vs. absence of midventral spines and presence of flat
surface venter of head and first trunk ring [Figs. 12–13, 15]).
Additional distinguishing characters include the absence of
bilateral bony lobed protuberances on the posterolateral
margins of the post-temporal bones (vs. presence in all
members of Acentronura and Idiotropiscis [Figs. 12, 16]), the
presence of the supracleithrum that extends dorsally from
the cleithrum, forming a continuous bony collar (Fig. 5) vs.
absence of a supracleithrum resulting in cleithra terminating
at the anterior margins of the bony dorsomedial crest [Figs.
10, 13, 14]), bony spines (on the snout, frontal, ventrolateral
surface of head, upper and lower cleithral ring, subdorsal
ridges, and principal trunk and tail ridges [Figs. 1–5] vs.
absence of spines except on the snout in I. australe and I.
larsonae [Figs. 12–13, 15]; four subdorsal spines, large cleithral
ring spine at level of last pectoral-fin ray, and small spine
present posterolaterally of the pelvic-fin base in I. lumnitzeri
[Figs. 13–15]). Cylix tupareomanaia can be further distin-
guished from the members of Idiotropiscis by the number of
subdorsal rings (3 vs. 4), tail rings (35–36 vs. 37 in I. australe,
39–40 in I. larsonae,43inI. lumnitzeri), and the tail rings
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Fig. 15. lCT scanned skeleton of Idiotropiscis lumnitzeri, CAS HH-0423, male, 69.9 mm SL. (A) Osteocranium in lateral view highlighting the ventral
cleithral cheek spine and small spine posteroventral of pectoral-fin base. (B) Subdorsal-fin base in lateral view highlighting the one upper and one
lower subdorsal spines. Abbreviations: CS, cleithral spine; LSDS, lower subdorsal spine; PLS, posterolateral spine on pectoral-fin base; USDS, upper
subdorsal spine.
Short and Trnski—New pygmy pipehorse from New Zealand 829
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Fig. 16. Lateral view of lCT scanned neurocranium of (A) Cylix tupareomanaia, NMNZ P.046322; (B) Hippocampus barbouri, USNM 220605.
Abbreviations: ANP, anterior nuchal plate; CO, coronet on anterior nuchal plate; PNP, posterior nuchal plate; SC, supracleithrum; SOC, supraoccipital.
Fig. 17. Phylogenetic hypothesis using nuclear gene sequences TMO-4C4 and 18S retrieved with Maximum Likelihood (ML), Maximum Parsimony
(MP), and Bayesian Inference (MrBayes), representing 17 species from clade 6 from the analysis of Hamilton et al. (2017) and the new taxon. Tree
rooted with the southern Australian trunk-brooder pipefish Heraldia nocturna. Nodal support at the generic level is shown in ML/MP/MrBayes order.
See Data Accessibility for tree file.
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spanning the male brood pouch (8 vs. 10 in I. australe,8inI.
larsonae,10inI. lumnitzeri [Table 4]).
The morphologically similar members of Acentronura (Figs.
8–9) share a number of characteristics of the head with Cylix
and Idiotropiscis, including: the absence of anterior nuchal
plate, presence of posterior nuchal plate, with large gap
between it and the supraoccipital; presence of the bony
dorsomedial crest on the posterior nuchal plate; and fully
enclosed male brood pouch enfolded by 7, 14, and 12 pouch
plates in A. breviperula,A. gracilissima, and A. tentaculata,
respectively (vs. 10 in C. tupareomanaia and 9–11 in Idio-
tropiscis [Table 4]). Acentronura can be distinguished by the
slim trunk (vs. deep trunk), continuous superior trunk ridge
(vs. discontinuous), low supraoccipital (vs. elevated supraoc-
cipital), distinct and low dorsomedial bony crest followed by
a raised bony dimple on the supraoccipital (vs. prominent
dorsomedial bony crest protruding anteriorly in Idiotropiscis;
and large cup-like pentamerous crest protruding anteriorly in
C. tupareomanaia), presence of bilateral bony lobed protuber-
ances on the posterolateral margins of the post-temporal
bones (vs. present in Idiotropiscis; absent in C. tupareomanaia);
absence of body spines (vs. snout spines in I. australe and I.
larsonae; four subdorsal spines, large cleithral spine at level of
last pectoral-fin ray, and small spine present posterolaterally
on the pelvic-fin base in I. lumnitzeri; and spines on snout,
frontal, ventrolaterally of head, upper and lower cleithral
ring, subdorsal ridges, and principal trunk and tail ridges in
C. tupareomanaia), number of tail rings (43 vs. 37 in I.
australe, 39–40 in I. larsonae,43inI. lumnitzeri; 35–36 in C.
tupareomanaia [Tables 3, 4]), and the number of tail rings
spanning the male brood pouch (14 vs. 11 in I. australe,9inI.
larsonae,11inI. lumnitzeri;8inC. tupareomanaia [Table 4]).
The anteriormost pouch plate is broad and paddle-shaped at
the ventrocaudal margin and curved posterolaterally relative
to the other plates, which are uniform in shape, in A.
gracilissima,A. tentaculata,I. australe, and I. lumnitzeri (vs. all
uniform in shape in C. tupareomanaia; unknown in A.
breviperula and I. larsonae). The brood pouch is further
distinguished in A. gracilissima by the first and second
anteriormost pouch plates, which appear to be large and
bifurcated ventrolateral extensions (Fig. 9B). Additionally,
the congener A. breviperula shares the fully enclosed male
brood pouch enfolded by arcuate ventrolateral bony exten-
sions (Fig. 12,Table 4).
Morphological comparisons to seahorses.—Cylix and Idio-
tropiscis are superficially similar in appearance to members
of Hippocampus and share with the latter numerous morpho-
logical synapomorphies, including meristic characters, an
elevated frontal and supraoccipital, principal trunk ridge
configuration, elevated dorsal-fin base, head angled ventrally
relative to the principal body axis (258vs. 908in seahorses),
moderately deep trunk (vs. slender trunk in Acentronura),
male brood pouch, prehensile tail, and absence of a caudal
fin. Cylix tupareomanaia, in particular, exhibits seahorse-like
characteristics, including head and principal body ridge
spines present on the snout, frontal, cleithral ring, and
subdorsal area, and a highly derived supraoccipital crest
present anterodorsally on its head that is remarkably similar
in outward appearance to the distinct coronet present in
most species of Hippocampus (Fig. 16, Table 5). In 2006, the
first photographic record of C. tupareomanaia, observed in
coralline and red algae at 10 m depth, was taken at Tawhiti
Rahi, the Poor Knights Islands, Northland, New Zealand
(Kuiter, 2009: 93, figs. A, B). However, the female individual
in the photograph was originally misidentified as the
subtropical Southwest Pacific seahorse, Hippocampus jugumus
Kuiter, 2001, due to a superficial similarity in external
appearance to this species (e.g., slender body, distinct
coronet, continuous cleithral ring, and meristic characters)
and therefore at the time mis-recorded as a range extension
from its type locality of Lord Howe Island, NSW, Australia to
temperate New Zealand. Hippocampus jugumus has since been
collected from New Zealand waters, at the offshore subtrop-
ical Rangit¯
ahua Kermadec Islands in the northernmost
region of New Zealand (Clark et al., 2017).
Despite the similarity between Cylix and Hippocampus,a
suite of anatomical features of the head and first trunk ring
(Fig. 16, Table 5) readily distinguish Cylix from Hippocampus,
including: anterior nuchal plate absent (vs. present in the
modified form of a coronet in Hippocampus between the
supraoccipital and the posterior nuchal plate); the elevated
supraoccipital well separated from the cleithrum above the
operculum, forming a large gap between the two areas (vs.
supraoccipital and anterior and posterior nuchal plates
interdigitated dorsally and connected lateroventrally by the
cleithrum); distinct coronet present but derived from the
cup-like and divided pentamerous bony crest on the
supraoccipital bone (vs. derived from the modified anterior
nuchal); presence of dorsomedial bony crest on the posterior
nuchal plate (vs. absence); cleithrum reaches mid-dorsally
over anterior margins of bony dorsomedial crest (vs. ventrally
on a projected horizontal line through gills in members of
Hippocampus with the exception of H. jugumus [Kuiter, 2001]
and the pygmy seahorse species H. japapigu,H. pontohi,H.
satomiae, and H. waleananus [Short et al., 2018]); snout spines
present on the mesethmoid and angled dorsally (vs. present
directly anterior of the interorbital space and angled
anteriorly); elevated rim of orbit protruding dorsally and
ventrally (vs. no elevation of rim of orbit); eye spines absent
(vs. presence of spines dorsally and ventrally of eye); strongly
elevated ventrolateral bulge on first trunk ring supporting the
pectoral fin (vs. absence of ventrolateral bulge on first trunk
ring); and male brood pouch plates present (vs. absent).
Genetic distances and phylogenetics.—Table 6 summarizes
uncorrected p-distances at the COX1 gene between C.
tupareomanaia and the previously sequenced pygmy pipe-
horses A. breviperula,A. tentaculata, and I. lumnitzeri (Ham-
ilton et al., 2017). Cylix tupareomanaia differs from A.
breviperula by 19.5%, A. tentaculata by 20.4%, I. australe by
14.1%, and I. lumnitzeri by 18.4%.
Genetic analyses performed here using the nuclear gene
sequences TMO-4C4 and 18S, and retrieved with ML, MP,
and MrBayes (Fig. 17), resulted in well-supported topologies
concordant with the hypothesized clade Vi of Hamilton et al.
(2017). The pygmy pipehorse genera Cylix,Idiotropiscis, and
Acentronura form a monophyletic group with strong support
in which C. tupareomanaia is placed as sister group to a clade
comprising Acentronura and Idiotropiscis. Together, the three
genera form the sister group to a clade comprising the Indo-
Pacific pipefish genera Filicampus,Haliichthys,Lissocampus,
and Trachyrhamphus, all of which are endemic or occur in
Australia and New Zealand. All of these aforementioned
genera together form a clade representing the sister group to
Hippocampus. The placement of C. tupareomanaia in clade Vi
Short and Trnski—New pygmy pipehorse from New Zealand 831
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further supports the refutation by Hamilton et al. (2017) of
the immediate sister taxon relationship between Hippocam-
pus and the pygmy pipehorses hypothesized by Teske and
Beheregaray (2009).
DATA ACCESSIBILITY
The tree file for the phylogenetic hypothesis associated with
this work (Fig. 17) is available at https://www.
ichthyologyandherpetology.org/i2020136. Unless an alterna-
tive copyright or statement noting that a figure is reprinted
from a previous source is noted in a figure caption, the
published images and illustrations in this article are licensed
by the American Society of Ichthyologists and Herpetologists
for use if the use includes a citation to the original source
(American Society of Ichthyologists and Herpetologists, the
DOI of the Ichthyology & Herpetology article, and any
individual image credits listed in the figure caption) in
accordance with the Creative Commons Attribution CC BY
License. ZooBank publication urn:lsid:zoobank.org:
pub:DD96034D-06EC-4FA5-9E68-B128195F88D3.
ACKNOWLEDGMENTS
We give sincere thanks to Hori Parata and kaum¯
atua of
Ng¯
atiwai for the partnership with the Auckland Museum and
in collaborating with the naming of Cylix tupareomanaia, and
for sharing their m¯
atauranga o Ng¯
atiwai. This collaboration
acknowledges Ng¯
atiwai as kaitiaki (guardians) of Cylix
tupareomanaia. We are grateful to Te Kahuratai Painting and
Daniel Hikuroa, University of Auckland, for the thoughtful
translation of the Abstract. Many individuals and institutions
contributed in the field, laboratory, observations, and
congenial discussions, and provided valuable specimens that
served as the basis for this study. We would like to
particularly thank Crispin and Irene Middleton for collecting
and photographing a paratype specimen from the Cavalli
Islands and the holotype at Waiatapaua Bay, Whangaruru;
Andrew Stewart, Museum of New Zealand Te Papa Tongar-
ewa, for providing access to the paratype specimen of C.
tupareomanaia; Severine Hannam, Auckland Museum, for
amazing curatorial assistance and collection efforts of the
holotype specimen in northern New Zealand; special thanks
to Kevin Conway, Texas A&M University, for editing the
manuscript; Oliver Crimmen and Ralf Britz, Natural History
Museum, London for assistance in providing x-ray radio-
graphs of the holotypes of A. breviperula and A. tentaculata;
David Catania, Jon Fong, and Mysi Hoang, California
Academy of Sciences, for amazing curatorial assistance; Mark
McGrouther, Amanda Hay, and Matt Lockett, Australian
Museum, for amazing curatorial assistance and specimen
loan of I. lumnitzeri; Ralph Foster, South Australian Museum,
for specimen loan of I. australe; Glenn Moore, Western
Australian Museum, for specimen loan of I. australe; Libby
Liggins, Institute of Natural and Mathematical Sciences,
Massey University, for providing COX1 sequences for C.
tupareomanaia and the nuclear sequences 18S and TMO-4C4
for I. australe; Avery Hiley and Greg Rouse, Scripps Institution
of Oceanography, UC San Diego, for providing the nuclear
sequences 18S and TMO-4C4 for C. tupareomanaia; Michael
Hammer, Museum and Art Gallery of the Northern Territory,
for photographs and x-rays of I. larsonae; Laith Jawad,
Freelance Fish Consultant, and Claire Attenborough for
providing x-rays of the type and comparative specimens;
Adam Summers and Jules Chabain, Friday Harbor Laborato-
ries, University of Washington, for micro-computed tomog-
raphy scans of pygmy pipehorse specimens used in this
study; Richard Smith, IUCN Seahorse, Pipefish and Seadra-
gon Specialist Group, Shane Housham and Julia Riddle,
Northland Dive, and Kent Erickson, Ocean Blue Adventures,
for high resolution underwater photographs of C. tupareo-
manaia; Alison Perkins, underwater photographer, for under-
water photographs and observations of C. tupareomanaia at
the type location; Carey Harmer, Leafy Seadragon Tours, for
many illuminating discussions on I. australe; Andrew Trevor-
Jones, Australian Museum, for many illuminating discussions
on I. lumnitzeri. Special thanks to David Harasti, Port
Stephens Fisheries Institute, Louw Classans, Rhodes Univer-
sity, and Maarten De Brauwer, Curtin University, for their
gushing support of all things syngnathids and lively
discussions; and Greg Misner, Aotearoa Hydrogen Alliance,
for the generation of the geographic information system
(GIS) map of the distribution of C. tupareomanaia within New
Zealand. Finally, we would like to express gratitude for the
work of C. E. Dawson, his classic 1985 book Indo-Pacific
Pipefishes: Red Sea to the Americas provided a comprehensive
assessment of syngnathid meristics and morphology for the
genera Acentronura and Idiotropiscis, which proved invaluable
for this study. The holotype and tail clipping from the type
locality were collected under Fisheries New Zealand Special
Permit number 691 issued to Auckland Museum. This
research was supported in part by funding from NSF (DBI-
1759637, DEB-170166) to Adam Summers, Friday Harbor
Laboratories, University of Washington.
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Appendix 1. GenBank numbers associated with mtDNA (COI) and nDNA (18S, TMO-4C4) sequences used in this study.
Species COI 18S TMO-4C4 Reference
Acentronura breviperula KY066067 KY065694 KY065608 Hamilton et al., 2017
Acentronura tentaculata KY066068 KY065695 KY065609 Hamilton et al., 2017
Cylix tupareomanaia, new species MH306206 MH286429 MH286430 This study
Filicampus tigris — KY065724 KY065634 Hamilton et al., 2017
Haliichthys taeniophorus — KY065729 KY065639 Hamilton et al., 2017
Heraldia nocturna — KY065730 KY065640 Hamilton et al., 2017
Hippocampus abdominalis — KY065731 KY065641 Hamilton et al., 2017
Hippocampus bargibanti — KY065732 KY065642 Hamilton et al., 2017
Hippocampus ingens — KY065734 KY065645 Hamilton et al., 2017
Hippocampus kuda — KY065733 KY065644 Hamilton et al., 2017
Hippocampus pontohi — KY065737 KY065648 Hamilton et al., 2017
Hippocampus reidi — KY065739 KY065650 Hamilton et al., 2017
Hippocampus trimaculatus — KY065740 KY065651 Hamilton et al., 2017
Hippocampus whitei — KY065741 KY065652 Hamilton et al., 2017
Hippocampus zosterae — KY065742 KY065653 Hamilton et al., 2017
Idiotropiscis australe MH306205 — — This study
Idiotropiscis lumnitzeri KY066120 KY065746 KY065657 Hamilton et al., 2017
Lissocampus caudalis — KY065749 KY065660 Hamilton et al., 2017
Lissocampus runa — KY065750 KY065661 Hamilton et al., 2017
Trachyrhamphus bicoarctatus — KY065781 KY065687 Hamilton et al., 2017
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