A New Genus and Species of the Frogfish Family Antennariidae (Teleostei: Lophiiformes: Antennarioidei) from New South Wales, Australia, with a Diagnosis and Key to the Genera of the Histiophryninae

Abstract

1 A new genus and species of the frogfish family Antennariidae, subfamily Histiophryninae, is described on the basis of three specimens collected near Kurnell and Bare Island in Botany Bay, New South Wales, Australia. It differs from all other antennariid genera in having a combination of features that includes a unique morphology of the first and second dorsal-fin spines, some or all fins fringed with red, and a unique combination of fin-ray and vertebral counts. The new genus is diagnosed, described, and compared to its sister genus, Kuiterichthys, using both molecular and morphological data. Notes on habitat and sponge mimicry, locomotion and defense, and reproduction and parental care are also provided as well as a diagnosis and a revised key to the known genera of the Histiophryninae. A N undescribed genus and species of the frogfish family Antennariidae (Lophiiformes, Antennarioi-dei), first collected on 15 October 1980 (UW 21020, 60 mm), was subsequently lost and additional specimens were not located again until 2005. Three specimens were subsequently collected on 30 October 2005, 18 January 2009, and 16 May 2009 by SCUBA divers near Kurnell and Bare Island in Botany Bay, New South Wales, Australia. Although frequently misidentified as a member of the genus Antennarius Daudin, 1816, by local divers, the new taxon more closely resembles the genus Kuiterichthys Pietsch, 1984. However, Kuiterichthys and the new genus are remarkably distinct, differing in a host of ways, the most striking of which is the morphology of the first and second dorsal-fin spines and distinctive red pigmentation on the distal margin of the fins (Figs. 1–3). The new species also has fewer anal-fin rays and a unique escal morphology.

Full text

A New Genus and Species of the Frogfish Family Antennariidae (Teleostei:
Lophiiformes: Antennarioidei) from New South Wales, Australia, with
a Diagnosis and Key to the Genera of the Histiophryninae
Rachel J. Arnold
1
, Rob Harcourt
2
, and Theodore W. Pietsch
1
A new genus and species of the frogfish family Antennariidae, subfamily Histiophryninae, is described on the basis of
three specimens collected near Kurnell and Bare Island in Botany Bay, New South Wales, Australia. It differs from all
other antennariid genera in having a combination of features that includes a unique morphology of the first and second
dorsal-fin spines, some or all fins fringed with red, and a unique combination of fin-ray and vertebral counts. The new
genus is diagnosed, described, and compared to its sister genus, Kuiterichthys, using both molecular and morphological
data. Notes on habitat and sponge mimicry, locomotion and defense, and reproduction and parental care are also
provided as well as a diagnosis and a revised key to the known genera of the Histiophryninae.
AN undescribed genus and species of the frogfish
family Antennariidae (Lophiiformes, Antennarioi-
dei), first collected on 15 October 1980 (UW 21020,
60 mm), was subsequently lost and additional specimens
were not located again until 2005. Three specimens were
subsequently collected on 30 October 2005, 18 January
2009, and 16 May 2009 by SCUBA divers near Kurnell and
Bare Island in Botany Bay, New South Wales, Australia.
Although frequently misidentified as a member of the genus
Antennarius Daudin, 1816, by local divers, the new taxon
more closely resembles the genus Kuiterichthys Pietsch, 1984.
However, Kuiterichthys and the new genus are remarkably
distinct, differing in a host of ways, the most striking of
which is the morphology of the first and second dorsal-fin
spines and distinctive red pigmentation on the distal margin
of the fins (Figs. 1–3). The new species also has fewer anal-
fin rays and a unique escal morphology.
MATERIALS AND METHODS
Standard length (SL) is used throughout. Methods for making
counts and measurements follow Pietsch and Grobecker
(1987). Terminology used to describe the various parts of the
angling apparatus follows Bradbury (1967). Symbolic codes
for institutions are those provided by Sabaj Pe´rez (2013).
Whole genomic DNA was extracted from skeletal muscle
of the holotype (UW 118988) and paratype (AMS I.44699)
of the new species using the QIAamp DNA Mini Kit
(Qiagen). Three outgroup taxa—the antennariids Echino-
phryne crassisspina McCulloch and Waite, 1918, Echino-
phryne mitchellii (Morton, 1897), and Phyllophryne scortea
(McCulloch and Waite, 1918)—were chosen based on
evidence from a previously published phylogeny (Arnold
and Pietsch, 2012). COI data from the second paratype
(AMS I.43749), Kuiterichthys furcipilis (Cuvier, 1817) (SAM
10476), and the outgroup taxa (NMV A25508.002, NMV
A29226, SAM 11544, SAM 11721) were also taken from the
study by Arnold and Pietsch (2012).
Approximately 674 base pairs of the 59region of the
cytochrome coxidase I (COI) gene were PCR amplified with
primer pairs COI5562f and COI6284r (Pietsch et al., 2009).
Thermocycle protocol followed the methods outlined by Ward
et al. (2005). Sequences were cleaned and generated at the
High-Throughput Genomics Unit, Department of Genome
Sciences, University of Washington, Seattle. All sequences
were aligned using MUSCLE (Edgar, 2004) as implemented in
GENEIOUS (Drummond et al., 2007). Sequence divergence
values and a neighbor-joining tree were generated with
GENEIOUS using the Tamura-Nei distance model. All new
sequences were deposited in GenBank (Table 1).
Porophryne, new genus
urn:lsid:zoobank.org:act:BAB46382-33B5-48DB-94C3-A11E
1333A2CA
Type species.—Porophryne erythrodactylus, new species
Diagnosis.—A member of the antennarioid family Antennar-
iidae, subfamily Histiophryninae (see below), distinguished
from its sister genus, Kuiterichthys, in having an entirely
different morphology of the second dorsal-fin spine (narrow
proximally, abruptly expanded laterally and becoming sub-
quadrangular in the new taxon vs. narrow and tapering
slightly toward the tip in Kuiterichthys) and esca (two tufts of
short filamentous appendages in the new taxon vs. many
filamentous appendages arising from a common base or
within a V-shaped depression in Kuiterichthys), and a
conspicuous tuft of filaments, usually brightly colored, on
the midline of the lip of the lower jaw that may serve as
accessory lure (unknown in any other anglerfish). It differs
further in having the following combination of character
states: all nine caudal-fin rays bifurcate, pectoral-fin rays 10 or
11 (8 on one side of smaller paratype), dorsal-fin rays 13, and
vertebrae 20 (determined by radiographs); some or all fins
fringed with red.
Etymology.—The generic name is derived from the Greek
poro, meaning to bear pores, in allusion to the naked, black,
‘‘osculum-like’’ spots present on the head and body of most
specimens, as well as its primary microhabitat within sponge
(Porifera) beds; and phryne, toad, a suffix commonly applied
to lophiiform generic names. The gender is feminine.
1
School of Aquatic and Fishery Sciences, and Burke Museum of Natural History and Culture, University of Washington, Campus Box 355020,
Seattle, Washington 98105-5020; E-mail: (RJA) schoenrj@uw.edu; and (TWP) twp@uw.edu.
2
Department of Biological Sciences, Macquarie University, North Ryde, Sydney, 2109, Australia.
Submitted: 2 December 2013. Accepted: 20 March 2014. Associate Editor: D. Buth.
F2014 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-13-155 Published online: October 21, 2014 ,
Copeia 2014, No. 3, 534–539

Porophryne erythrodactylus, new species
urn:lsid:zoobank.org:act:8C14ADFE-26E2-499F-BEDC-2F6E
25EDFB0C
Figures 1–3, Table 1
Antennariidae gen. et sp. nov.—Arnold and Pietsch, 2012:117,
124, 125, 128, figs. 1G, 5, 7, table 1 [molecular phylogeny].
Holotype.—Female, UW 118988, 54.6 mm, Australia, New
South Wales, Botany Bay, Kurnell, SCUBA, 8 m, D. Harasti,
16 May 2009.
Paratypes.—Two females: AMS I.43749, 68.3 mm, Australia,
New South Wales, Botany Bay, Bare Island, approximately
33u599S, 151u139E, SCUBA, 13.7 m, R. Harcourt, 30 October
2005; AMS I.44699, 74.3 mm, east side of Bare Island,
33u599330S, 151u139570E, 10.7 m, R. Harcourt, 18 January 2009.
Diagnosis.—As given for the genus.
Description.—Esca red, gray, or brown, divided into two tufts
of short filamentous appendages; first dorsal-fin spine
(illicium) 6.3–7.7%SL, sparsely covered with dermal denti-
cles, without groove alongside base of second dorsal spine;
second dorsal-fin spine 17.6–21.0%SL, without posterior
membrane, sometimes with slender filamentous appendages
distally, narrow for approximately one third of its length,
then abruptly expanded laterally and becoming sub-qua-
drangular; third dorsal-fin spine 27.4–35.3%SL, mobile,
covered with thick skin; diameter of eyes 6.9–7.5%SL;
dorsal-fin rays 13 (all bifurcate except anteriormost 1 or 2);
anal-fin rays 7 (all bifurcate); pectoral-fin rays 10 or 11 (8 on
left side of smaller paratype; all simple); pelvic-fin rays 5
(posteriormost ray bifurcate); caudal-fin rays 9 (all bifurcate).
Body everywhere covered with dermal denticles, except
within small black spots (see below); orange, yellow, pink,
red, and/or white individuals usually with brown and red
cutaneous appendages occurring in patches on body, espe-
cially around chin and opercle; a conspicuous cluster of
fleshy filaments, often brightly pigmented, on midline of
lip of lower jaw (possibly serving as an additional luring
device; Fig. 3E); oral valve present, lining both upper and
lower jaws; swimbladder present; ovaries paired, lobular;
caudal peduncle present, posteriormost rays of dorsal and
anal fins not connected by membrane to caudal peduncle
and outermost rays of caudal fin.
In life, pale to dark gray, orange and red, orange red and
pink, white and pink, white and red, or yellow and red, with
some or all fins fringed with red; first and second dorsal-fin
spines, premaxilla, and dentary with some red or matching
body coloration; tiny light to dark gray spots on gray
individuals, only visible on close inspection, giving the fish
an overall light to dark gray color, with some individuals
appearing nearly white; gray individuals usually with a
variable number of naked, black, osculum-like spots scat-
tered asymmetrically on head and body.
In preservation, dark to pale gray, fading ventrally, with
larger diffuse gray spots (covered with dermal denticles) on
abdomen; white where red pigment has leached away;
pigment of orange and red paratype (AMS I.44699) leached
away (except for margins that were red in life, now light
brown), patches of cutaneous appendages brown.
Molecular evidence.—An 8.4–9.5%divergence in the COI
gene was found between Phyllophryne and Echinophryne, two
well-established genera of the Antennariidae (Pietsch, 1984;
Pietsch and Grobecker, 1987; Arnold and Pietsch, 2012).
Between Porophryne and Kuiterichthys, 10.9–11.3%diver-
gence was calculated, and 100%identity was found between
the three individuals of Porophryne erythrodactylus.
Etymology.—The specific name erythrodactylus is derived
from the Greek erythros, meaning red; and dactylus, fingers,
in reference to the conspicuous red pigmentation on the tips
of the pectoral-fin rays as well as along the distal margins of
some or all other fins (Fig. 3B).
Geographic distribution.—The type material of Porophryne
erythrodactylus is known only from nearshore waters of
Botany Bay, New South Wales, at depths of 8–13.7 m, but
observations have been recorded in as little as 6 m and as
deep as 24 m (see Supporting Online Material). Most un-
collected specimens of P. erythrodactylus have been reported
from the Sydney region, especially around Bare Island in
Botany Bay, but confirmed observations have also been made
as far south as Jervis Bay, New South Wales (Fig. 2B).
Habitat and sponge mimicry.—Porophryne erythrodactylus is
generally found in subtidal rocky-reef habitat dominated by
small filamentous and foliose algae of the genera Zonaria,
Corallina,Amphiroa, and Laurencia (often with some Sargas-
sum spp.). A few scattered larger seaweeds, including
Ecklonia radiata and Sargassum spp., with large patches of
filamentous or foliose algae, may also be present (Creese et
Fig. 1. Porophryne erythrodactylus, new genus and species, holotype,
UW 118988, Kurnell, Botany Bay, NSW (photo by D. Harasti).
Arnold et al.—New genus and species of frogfish from Australia 535

Fig. 2. Porophryne erythrodactylus, new genus and species: (A) paratype, AMS I.44699, Bare Island, Botany Bay, NSW (photo by R. Harcourt); (B)
orange and red color morph, Middle Ground, Jervis Bay, NSW (photo by K. Sebo); (C) pink color morph, Bare Island, Botany Bay, NSW (photo by N.
Missenden); (D) gray color morph, Bass Point, Shellharbour, NSW (photo by M. McKnight); (E) pink and white color morph, Bass Point, Shellharbour,
NSW (photo by M. McKnight); (F) red, orange, and pink color morph, Bass Point, Shellharbour, NSW (photo by M. McKnight).
536 Copeia 2014, No. 3

Fig. 3. Porophryne erythrodactylus, new genus and species: (A) red and gray color morphs, Bare Island, Botany Bay, NSW (photo by D. Harasti); (B)
gray color morph, Kurnell, Botany Bay, NSW (photo by R. Arnold); (C) yellow color morph, Bare Island, Botany Bay, NSW (photo by N. Missenden);
(D) close-up of the second dorsal-fin spine and illicium and esca, Bare Island, Botany Bay, NSW (photo by N. Missenden); (E) close-up of tuft of
filaments on lower jaw that possibly serves as an additional luring device, Bare Island, Botany Bay, NSW (photo by S. Bowtle); (F) guarding a cluster of
eggs, Bass Point, Shellharbour, NSW (photo by M. McKnight).
Arnold et al.—New genus and species of frogfish from Australia 537

al., 2009). Vertical or sloping walls on the deep edge of
nearby reefs are inhabited by ascidians, corals, and sponges,
the latter including Spongia sp., Tedania anhelans,Ephydatia
fluviatilis,Darwinella australiensis,Chondrilla australiensis,
Mycale australis,andHolopsamma laminaefavosa.More
specifically, the new species is most often found closely
associated with small sponges, blending in with the gray
coloration of Psammocinia sp. or the red or yellow coloration
of Darwinella sp., or nestled next to fan-shaped Echinocla-
thria leporina. In addition, the black ‘‘osculum-like’’ spots
typically present on gray color morphs of the new species
presumably aid in the mimicry of sponges as well.
DISCUSSION
Porophryne erythrodactylus appears to have two distinct color
phases: gray individuals usually with naked black spots
scattered asymmetrically on the head and body, and a near
absence of cutaneous appendages on the body; and orange,
yellow, pink, red, and/or white individuals, typically
without naked black spots, but usually with many more
cutaneous appendages on the body, the latter presumably to
aid in camouflage or to enhance luring through mimicry of
the substrate (Figs. 2A, 2B, 2F, 3A, 3D). Orange, yellow, pink,
red, and/or white individuals are also usually larger than
gray individuals: of the three type specimens, the orange
and red paratype (AMS I.44699) is the largest at 74.3 mm
(Fig. 2A), while in situ photographs of red individuals found
near gray individuals are generally larger (Fig. 3A). We
initially speculated that P. erythrodactylus might be sexually
dichromic or that there were two undescribed species, but
dissections of the three specimens confirmed that all are
female, and divergence within the COI gene is 0%.
Locomotion and defense.—Unlike Histiophryne psychedelica,
Porophryne erythrodactylus has never been observed to fall
from elevated substrates and then push off objects or the
seafloor while it ‘‘hops’’ from one location to another
(Pietsch et al., 2009). It has also never been seen to use jet-
propulsion in a sustained, long-distance swimming effort
(Pietsch and Grobecker, 1987). Video recorded by S. Bowtle
and observations made by one of us (RJA) indicate that P.
erythrodactylus, instead, darts quickly over relatively short
distances, primarily by caudal locomotion. However, like
Histiophryne and other genera (e.g., Echinophryne and Phyllo-
phryne) observed by RJA, P. erythrodactylus often keeps its
caudal fin strongly curled to one side of the body. In another
video taken by S. Bowtle, in which the same individual
received some amount of prodding by the diver, the
individual eventually quit trying to swim away and, instead,
kept its tail strongly curled and fins and dorsal-fin spines
splayed. The frogfish maintained this position even when
picked up and dropped several times by the diver (see
Supporting Online Material). No member of the order
Lophiiformes has ever been shown to be toxic or venomous;
we therefore speculate that this behavior is its only defense
when camouflage fails—playing dead while making itself
appear as large as possible.
Reproduction.—Like other members of the subfamily Histio-
phryninae, the new species has lobular ovaries (Arnold and
Pietsch, 2012). Members of the Histiophryninae care for
their relatively small number of large eggs in various ways:
attached to the body, as in Lophiocharon (Pietsch and
Grobecker, 1980:figs. 1, 2, as Antennarius caudimaculatus;
Arnold and Pietsch, 2012:fig. 2D); carried within a pocket
formed by the anal, caudal, and dorsal fins, as in Histiophryne
(Pietsch and Grobecker, 1987:pl. 53; Pietsch et al., 2009:fig.
8); and nest guarded, as in Rhycherus (Kuiter, 1993:49,
unnumbered fig.). The offspring undergo direct develop-
ment, hatching as benthic juveniles, and without post-
hatching parental care. On 29 October 2011, an individual
of Porophryne erythrodactylus (not collected) was discovered
guarding a clutch of relatively large eggs at Bass Point,
Shellharbour, New South Wales (Fig. 3F), confirming that
the new species is a nest-guarder. Unfortunately, neither the
sex of the individual guarding the eggs nor the number of
eggs could be determined from the photograph.
DIAGNOSIS OF THE HISTIOPHRYNINAE
Histiophryninae Arnold and Pietsch, 2012
Histiophryninae.—Arnold and Pietsch, 2012:117, 124–128,
figs. 5, 7 [molecular phylogeny]; Arnold, 2013:496
[additional species].
Type genus.—Histiophryne Gill, 1863, by original description.
Diagnosis.—Histiophryninae, a subfamily of the Antennar-
iidae, distinguished from its sister group, the Antennariinae,
in having lost the mesopterygoid and epural (Pietsch, 1984;
Pietsch and Grobecker, 1987). The two subfamilies differ
also in having an entirely different ovarian morphology: the
Histiophryninae has simple oval-shaped ovaries, while the
Antennariinae has double scroll-shaped ovaries (Pietsch and
Grobecker, 1987:pl. 10, fig. 161); each ovarian type corresponds
to a different life history: members of the Histiophryninae
undergo direct development and display various degrees of
parental care, while those of the Antennariinae are broadcast
spawners and go through a distinct larval stage. In addition,
the Histiophryninae is restricted geographically to the Indo-
Australian Archipelago, while the Antennariinae has a broad
geographic distribution, with all known genera found cir-
cumglobally throughout the tropics and subtropics.
KEY TO THE KNOWN GENERA OF THE HISTIOPHRYNINAE
1a. Second and third dorsal-fin spines hidden, laid back
and bound to surface of cranium by skin of head;
caudal peduncle absent, membranous posterior-
most margin of soft dorsal and anal fins extending
posteriorly beyond base of caudal fin and connect-
ing to outermost caudal-fin rays
__________________________________ _____
_________________________________ ____________________________
Histiophryne Gill, 1863
Table 1. Tissue and voucher specimens of antennariids with
corresponding GenBank accession number for the cytochrome
oxidase csubunit 1 (COI) gene.
Species Voucher GenBank no.
Echinophryne crassispina SAM 11544 GU188510
Echinophryne mitchellii NMV A25508.002 FJ224368
Kuiterichthys furcipilis SAM 10476 GU188504
Porophryne erythrodactylus UW 118988 KF802202
Porophryne erythrodactylus AMS I.43749 GU188506
Porophryne erythrodactylus AMS I.44699 KF802201
Phyllophryne scortea NMV A29226 GU188509
Phyllophryne scortea SAM 11721 FJ224369
538 Copeia 2014, No. 3

1b. Second and third dorsal-fin spines free, not hidden
beneath skin of head; caudal peduncle present or
absent, dorsal and anal fins terminating well before
or at junction of caudal fin, but not broadly
attached to it
____________________________________________________________________________
2
2a. All rays of caudal fin simple
____________________________________________
____________________________________________________
Tathicarpus Ogilby, 1907
2b. All or at least seven innermost caudal-fin rays
bifurcate
______________________________________________________________________________________
3
3a. All caudal-fin rays bifurcate
_______________________________________________
4
3b. Seven innermost caudal-fin rays bifurcate, outer-
most rays simple
_____________________________________________________________________
7
4a. Skin smooth, without dermal denticles, but cov-
ered with elongate, close-set cutaneous appendag-
es; third dorsal-fin spine free, only proximal 20–
25%connected to nape of neck by membrane
______
_______________________________________________________
Rhycherus Ogilby, 1907
4b. Skin covered with dermal denticles; third dorsal-fin
spine connected for more than 25%of its length to
nape of neck by membrane or connected to soft
dorsal fin by skin
____________________________________________________________________
5
5a. Second dorsal-fin spine long, greater than 17%SL,
expanded laterally after approximately one third its
length, becoming sub-quadrangular; naked black
spots often present on head and body; distal tip of
some or all fin rays red
____________
Porophryne, new genus
5b. Second dorsal fin short (less than 9%SL) or
connected to base of third spine, never narrow at
base and expanded distally; head and body with or
without black spots, but spots never naked; distal
tip of fin rays only rarely red
____________________________________________
6
6a. Second dorsal-fin spine connected to base of third
spine, third spine connected to soft dorsal fin by
skin; caudal fin almost always with translucent
ocelli between each two rays
__________________________________________
_____________________________________________
Lophiocharon Whitley, 1933
6b. Second dorsal-fin spine free from third spine, third
dorsal-fin spine not connected to soft dorsal fin by
skin; caudal fin without translucent ocelli between
rays
__________________________________________
Allenichthys Pietsch, 1984
7a. Skin smooth, without dermal denticles; caudal
peduncle absent, membranous posteriormost mar-
gin of soft dorsal and anal fins attached to body at
base of outermost rays of caudal fin
____________________________
________________________________________________
Phyllophryne Pietsch, 1984
7b. Skin rough, covered with close-set dermal denti-
cles; caudal peduncle present, membranous poste-
riormost margin of soft dorsal and anal fins
attached to body distinctly anterior to base of
caudal fin
____________________________________________________________________________________
8
8a. Illicium completely covered with close-set dermal
denticles; bulbous esca absent
________________________________________
_______________
Echinophryne McCulloch and Waite, 1918
8b. Illicium naked, without dermal denticles; esca dis-
tinct, containing several filamentous appendages
______
______________________ _________________________________
Kuiterichthys Pietsch, 1984
SUPPORTING ONLINE MATERIAL
Additional records and in-situ photographs of Porophryne
erythrodactylus posted on the Australian Museum website:
http://australianmuseum.net.au/Undescribed-Anglerfish
In-situ videos of Porophryne erythrodactylus: in surrounding
habitat (courtesy of N. Missenden): http://www.youtube.
com/watch?v59sduMmK6pfo; swimming (courtesy of S.
Bowtle): http://www.youtube.com/watch?v5dREVud3Av8s;
playing dead (courtesy of S. Bowtle): http://www.youtube.
com/watch?v5x1M-1sm0YxA
ACKNOWLEDGMENTS
We thank the following individuals and their institutions for
providing loans and collection data: A. Hay, M. McGrouther,
and S. Reader (AMS), and K. Maslenikov (UW). We thank the
following individuals for the use of their photographs and
videos: S. Bowtle, D. Harasti, M. McKnight, N. Missenden, and
K. Sebo. We especially thank the following individuals for
collecting specimens: J. Coombs, R. Fea, D. Harasti, and D.
Ross. The molecular work was carried out in the Molecular
Ecology Research Laboratory, School of Aquatic and Fishery
Sciences, University of Washington. This work was supported
by the Dorothy T. Gilbert Ichthyology Research Fund through
the School of Aquatic and Fishery Sciences, University of
Washington.
LITERATURE CITED
Arnold,R.J.2013. A new species of frogfish of the genus
Kuiterichthys (Lophiiformes: Antennariidae: Histiophryninae)
from New South Wales, Australia. Zootaxa 3718:496–499.
Arnold, R. J., and T. W. Pietsch. 2012. Evolutionary history
of frogfishes (Teleostei: Lophiiformes: Antennariidae): a
molecular approach. Molecular Phylogenetics and Evolu-
tion 62:117–129.
Bradbury, M. G. 1967. The genera of batfishes (family
Ogcocephalidae). Copeia 1967:399–422.
Creese, R. G., T. M. Glasby, G. West, and C. Gallen. 2009.
Mapping the habitats of NSW estuaries. Industry and
Investment NSW Fisheries Final Report Series 113. Nelson
Bay, NSW, Australia. ISSN 1837-2112.
Drummond, A. J., M. Kearse, J. Heled, R. Moir, T. Thierer, B.
Ashton, A. Wilson, and S. Stones-Havas. 2007. GEN-
EIOUS, v. 3.7. Biomatters Ltd., Auckland, New Zealand.
Edgar, R. C. 2004. MUSCLE: multiple sequence alignment
with high accuracy and high throughput. Nucleic Acids
Research 32:1792–1797.
Kuiter, R. H. 1993. Coastal Fishes of South-Eastern Australia.
University of Hawaii Press, Honolulu, Hawaii.
Pietsch, T. W. 1984. The genera of frogfishes (family
Antennariidae). Copeia 1984:27–44.
Pietsch, T. W., R. J. Arnold, and D. J. Hall. 2009. A bizarre
new species of frogfish of the genus Histiophryne (Lophii-
formes: Antennariidae) from Ambon and Bali, Indonesia.
Copeia 2009:37–45.
Pietsch, T. W., and D. B. Grobecker. 1980. Parental care as
an alternative reproductive mode in an antennariid
anglerfish. Copeia 1980:551–553.
Pietsch, T. W., and D. B. Grobecker. 1987. Frogfishes of the
World: Systematics, Zoogeography, and Behavioral Ecol-
ogy. Stanford University Press, Stanford, California.
Sabaj Pe´rez, M. H. (Ed.). 2013. Standard symbolic codes for
institutional resource collections in herpetology and
ichthyology: an online reference. Version 4.0 (28 June
2013). American Society of Ichthyologists and Herpetol-
ogists, Washington, D.C. Electronically accessible at
http://www.asih.org
Ward, R. D., T. S. Zemlak, B. H. Innes, P. R. Last, and
P. D. N. Hebert. 2005. DNA barcoding Australia’s fish
species. Philosophical Transactions of the Royal Society B
360:1847–1857.
Arnold et al.—New genus and species of frogfish from Australia 539