RECORDS OF THE
AUSTRALIAN MUSEUM
Volume 67
Number 7 2 December 2015
Taxonomic Resolution to the Problem of
Polyphyly in the New Caledonian Scincid Lizard Genus
Lioscincus (Squamata: Scincidae)
by
Ross A. Sadlier, Aaron M. Bauer, Glenn M. Shea, and Sarah A. Smith
Table of Contents 2013-2015
Editorial Board
Dr Shane Ahyong
Dr Don Colgan
Dr Elena Kupriyanova
Dr Andrew Mitchell
Dr Robin Torrence
Dr Ross Sadlier
Editor
Dr Shane McEvey
Journal compilation © 2015 Australian Museum, Sydney
No part of this publication may be reproduced without
permission of the Editor.
Volume 67 Number 7
Published (print and online) 2 December 2015
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© The Authors, 2015. Journal compilation © Australian Museum, Sydney, 2015
Records of the Australian Museum (2015) Vol. 67, issue number 7, pp. 207-224.
ISSN 0067-1975 (print), ISSN 2201-4349 (online)
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1649
Taxonomic Resolution to the Problem of Polyphyly in
the New Caledonian Scincid Lizard Genus Lioscincus
(Squamata: Scincidae)
Ross A. Sadlier 1 *, Aaron M. Bauer 2 , Glenn M. Shea 3 ’ 1 and Sarah A. Smith 1
1 Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney NSW 2010, Australia
2 Department of Biology, Villanova University,
800 Lancaster Avenue, Villanova, Pennsylvania 19085, United States of America
3 Faculty of Veterinary Science B01, University of Sydney NSW 2006, Australia
Abstract. Recent genetic studies have identified the New Caledonian scincid genus Lioscincus to be
polyphyletic, comprising four distinct evolutionary lineages which we recognize at the generic level. The
revised concept of Lioscincus s.s. now includes only the type species Lioscincus steindachneri Bocage,
1873 and the recently described Lioscincus vivae Sadlier, Bauer, Whitaker & Smith, 2004. The three
remaining lineages identified are: Leiolopisma tillieri Ineich & Sadlier, 1991 and Lioscincus maruia Sadlier,
Whitaker & Bauer, 1998 for which the genus Phasmasaurus gen. nov. is proposed; Lygosoma (Mocoa)
nigrofasciolatus Peters, 1869 and Leiolopisma greeri Bohme, 1979 for which the genus Epibator gen. nov.
is proposed; and Lygosoma {Leiolopisma) novaecaledoniae Parker, 1926 for which the genus Caesoris
gen. nov. is proposed. Each of these genera is diagnosed by a suite of morphological apomorphies which
in combination is unique within the Eugongylus group of skinks of which each is a member. The revised
taxonomy presented here, which recognises an additional three new genera, brings the number of endemic
skink genera present in New Caledonia to 17, representing approximately 40% of the generic diversity
within the Australian/Pacific region Eugongylus group of skinks as currently recognised. Although we
are reluctant to erect new genera to accommodate one or two species, the data at hand suggest this as the
most accurate reflection of the available genetic and morphological data.
Keywords. Taxonomy; scincid; new genera; Lioscincus ; New Caledonia.
Sadlier, Ross A., Aaron M. Bauer, Glenn M. Shea, and Sarah A. Smith. 2015. Taxonomic resolution to the
problem of polyphyly in the New Caledonian scincid lizard genus Lioscincus (Squamata: Scincidae). Records of the
Australian Museum 67(7): 207-224.
The discovery and description of the New Caledonian
scincid lizard fauna covers a period of just over 150 years.
The first century of investigation in the period 1869-1970
saw contributions by five workers (Bavay, 1869; Peters,
1869, 1879; Gunther, 1872; Bocage, 1873; Brocchi, 1876)
describing 17 species in the nineteenth century, and six
* author for correspondence
workers (Andersson, 1908; Werner, 1909; Roux, 1913;
Parker, 1926; Mertens, 1928; Loveridge, 1941) describing
nine species or subspecies in the early twentieth century.
Twenty of these 26 taxa remain valid today. The most
significant contributions were the Catalogue des reptiles de
laNouvelle-Caledonie et description d’especes nouvelles of
208
Records of the Australian Museum (2015) Vol. 67
Bavay published in 1869 and the monograph Les reptiles de
la Nouvelle-Caledonie et des ties Loyalty of Roux published
in 1913. Bavay’s work described seven of the species
recognized today, while Roux provided a synthesis of his
own discoveries and observations with the contributions
of the previous 40 years. The majority of New Caledonian
species recognized by Roux were included within four of the
11 subgenera that resided under an expansive Lygosoma, at
that time one of the most diverse genera in terms of number
of species and worldwide distribution.
During this first century of research, generic conservatism
in skink systematics was a consistent theme. The recognition
of increasingly finer subgeneric units by Smith (1937) and
Mittleman (1952) began the process of recognizing groups
of species that later formed the basis of many of the skink
genera recognized today. However, as classifications that
reflected natural groupings, particularly with regard to
the New Caledonian species, these were only a marginal
improvement over earlier taxonomies. As such, the
systematic status of the lineages of “lygosomid” skinks,
the group in which the majority of New Caledonian skink
fauna was placed, remained largely unresolved for over 100
years, with the consequence that the generic diversity of the
island’s skink fauna was grossly under-appreciated (Bauer,
1989; Bauer & Sadlier, 2000), as was that of other regional
skink faunas belonging to this group.
In the latter part of the twentieth century, Allen Greer
began a 35 year refinement of scincid systematics which re¬
defined the subfamily Lygosominae (Greer, 1970), identified
major subgroups within this subfamily (Greer, 1974, 1979,
1989), and extensively reviewed lygosomine genera and
their relationships, identifying and refining characters for
diagnosing natural groups in the process. The contributions of
Greer had a major impact on understanding the relationships
of the Australian and Pacific region skink fauna, including
the establishment over time of a systematic framework
within which the intra- and inter-generic relationships of
this fauna could be assessed (see below for a more detailed
account of Greer’s contributions, and the implications of his
work on defining lineages of New Caledonian skinks and the
relationships of these lineages to other skinks).
It is within the context of this framework that investiga¬
tions into the diversity and relationships of the New
Caledonian skink fauna have been conducted over much of
the past 30 years, considerably changing previous concepts
of the species richness and generic diversity for this regional
fauna. During this period the recognition and definition of
monophyletic groups within the New Caledonian skinks has
largely been undertaken within the phylogenetic framework
for the Eugongylus group established by Greer (1974,
1979, 1989), and utilized similar operational principles
in identifying suites of morphological synapomorphies to
define natural or monophyletic genera (Sadlier, 1987; Sadlier
& Bauer, 1997, 1999b, 2000, 2002a; Sadlier et al. , 1997,
2004b, 2006b), and intrageneric groupings (Sadlier, 1990;
Sadlier et al ., 2002b).
Later studies, the more recent of which have utilized
morphological data in combination with genetic information
derived from DNA sequence data, have further identified and
defined supraspecific groups of New Caledonian skink taxa
as distinct evolutionary lineages (Smith et al., 2007; Ineich
et al., 2014), in some cases drawing extensively from the
genetic data for support for monophyly of the constituent
species (Sadlier et al., 2004a), and as support for taxa with
limited morphological apomorphies as discrete evolutionary
entities (Sadlier et al., 2014b). As a result the diversity in
terms of the number of skink genera strictly endemic to
New Caledonia has nearly doubled, with the recognition
of an additional six genera since 1987. Contributing to this
increased island generic endemism has been the transfer
of the New Caledonian species previously retained in
Leiolopisma to Lioscincus (Bauer & Sadlier, 1993), a
redefined Nannoscincus that includes only New Caledonian
taxa (Sadlier et al, 2006a) and five newly erected genera
(Lacertoides Sadlier, Shea & Bauer; Simiscincus Sadlier &
Bauer; Kanakysaurus Sadlier, Smith, Bauer & Whitaker;
Celatiscinciis Sadlier, Smith & Bauer; Phaeoscincus Sadlier,
Bauer, Smith, Shea & Whitaker) to accommodate new taxa.
With these additions there are currently 16 genera of skinks
in New Caledonia, of which 14 are either strictly endemic,
or nearly so, to the region, with more than 60 described
constituent species, over half of which have been discovered
and described in the past quarter of a century.
These earlier studies that defined natural groups on
morphological characters provided the platform for a
comprehensive genetic analysis of the intra and intergeneric
relationships of the New Caledonian taxa, and of these with
other members of the Eugongylus group of skinks (Smith et
al, 2007). This study identified the endemic New Caledonian
genera, together with the New Zealand skink radiation,
as a monophyletic lineage restricted to the Gondwanan
Tasman continental block. It also recognized a number of
suprageneric groups within the New Caledonian radiation,
and identified the taxa within Lioscincus as belonging to
four independent unrelated lineages, which are the focus
of the investigation presented here. A later genetic study
concentrating on the endemic New Zealand skink fauna
(Chappie et al, 2009), but using a larger number of genes,
confirmed the endemic New Caledonian and New Zealand
skink genera as a monophyletic lineage with the New
Caledonian and New Zealand skink fauna as discreet sister
lineages, and also the polyphyly of Lioscincus. Further
contributions highlighting the polyphyly of Lioscincus were
apparent in the molecular phytogenies of Caut et al. (2013)
and Ineich et al. (2014) in investigating the relationships
of the giant New Caledonian skink Phoboscincus bocourti
(Brocchi). It is the purpose of this paper to diagnose the
evolutionary lineages currently included under Lioscincus
and to erect new genera for those taxa no longer included
in that genus.
Leiolopisma and its role in
New Caledonian skink systematics
Greer’s early work on lygosomine skinks identified the
“greatest single taxonomic problem with the Lygosominae”
as the “delimitation and relationships of genera” (Greer,
1970, p. 171), and in particular the relationships within
“Leiolopisma” “as a major unresolved problem in skink
systematics” (Greer, 1970, p. 179). Generic concepts
were refined by Greer (1974), who diagnosed a number of
monophyletic genera from Malcolm Smith’s (1937) Section
Leiolopisma, and allocated these and related genera to three
major groups based on a suite of osteological and scalation
characters. Within this scheme of relationships, the New
Caledonian skink species known at the time all resided in
Sadlier et al .: New Caledonian lizard genus Lioscincus
209
genera placed in a large group of taxa, later formally referred
to as the “ Eugongylus Group” (Greer, 1979). The majority of
the New Caledonian species were retained within a redefined
Leiolopisma, which also included species from New Zealand,
Australia (including Lord Howe Island), and Mauritius in the
Indian Ocean. Greer (1974, p. 17) seems to have regarded
Leiolopisma as a (monophyletic) lineage. However, the genus
was weakly diagnosed, with exceptions amongst the included
species for almost every diagnostic character presented. The
significance of Greer’s early contributions to the systematics
of the New Caledonian skinks lies with being the first worker
to assess relationships and diagnose natural groups among
related lygosomine skinks in a contemporary phylogenetic
context, and in doing so provided a considerably more
robust framework within which to investigate interspecific
and intergeneric relationships of the New Caledonian taxa.
The major groups of genera identified by Greer within
the Lygosominae have proven remarkably robust, and
subsequent studies (Austin & Arnold, 2006; Brandley etal.,
2005; Honda etal., 2000,2003; Pyron etal., 2013; Rabosky
et al., 2007; Skinner et al., 2011; Whiting et al., 2003) have
largely supported these groupings.
Field studies initiated in 1978 were the basis of a
monograph of the scincid lizards by Sadlier (1987), the first
review in over 70 years, which recognized a doubling in
the number of endemic species to 24 and a similar increase
in the number of genera from 4 to 10, of which 7 were
strictly endemic. Sadlier (1987) proposed several putatively
monophyletic genera ( Caledoniscincus, Marmorosphax
and Sigaloseps, and a resurrected Tropidoscincus Bocage,
1873) to accommodate most of the New Caledonian species
formerly assigned to Leiolopisma. All were defined by
suites of apomorphic morphological character states, the
polarity of which was determined within the context of
Greer’s (1979) Eugongylus group of skinks. However,
four species with relatively plesiomorphic morphology
(nigrofasciolatus Peters, greeri Bohme, steindachneri
Bocage and novaecaledoniae Parker) were retained within
the “cosmopolitan” Leiolopisma, though two of these
{nigrofasciolatus and greeri) were recognized at the time as
probable sister taxa in a putative sublineage. Subsequently
three more New Caledonian species were added to the
genus, Leiolopisma tillieri Ineich & Sadlier in 1990, maruia
Sadlier, Whitaker & Bauer in 1998 (by which time the New
Caledonian Leiolopisma had been moved to Lioscincus,
see below), and vivae Sadlier, Bauer, Whitaker & Smith (as
Lioscincus) in 2004.
The members of “Leiolopisma” at the time of Sadlier’s
monograph (1987) also included all New Zealand skinks (20
species), a group of Australian skinks (13 species that now
represent Bassiana, Pseudemoia in part, and Niveoscincus
in part), the Lord Howe/Norfolk Island species lichenigera,
a single Fijian species {alazon Zug), and the type species
for the genus, Leiolopisma telfairii from Mauritius. The
diagnosis for “ Leiolopisma ” given by Greer (1970) contained
both “primitive” and “derived” character traits with both
states present for some characters (frontoparietal scales),
but was later modified by Sadlier (1987) to include only
the single most unifying apomorphy to define the genus,
lack of supranasal scales in most species. Further, limited
knowledge of morphological variation for some characters
and the simplistic assumption that some states, such as the
presence of supranasal scales and presence of a scaled lower
eyelid (both considered plesiomorphic) were homologous
across genera, would ultimately prove to be misleading, and
clouded interpretations of relationships within this group of
species well into the future. In effect “ Leiolopisma ” at this
time remained (as before) a genus of convenience rather
than one reflecting defined evolutionary relationships, and
the New Caledonian taxa retained within it (Sadlier, 1987)
were the residual species lacking a distinct supranasal scale
that could not be otherwise accommodated within any of
the existing putatively monophyletic genera within the
Eugongylus group.
A decade after Greer defined the Eugongylus group, two
not entirely concordant works were published which were
to influence generic concepts of this group of skinks in
the years following, and in particular the dynamics of taxa
formerly included in the all-encompassing “ Leiolopisma ” of
earlier studies. In one Greer (1989) diagnosed a Pseudemoia
subgroup within the Eugongylus group on the basis of
a single apomorphy not observed in other major groups
of skinks, fusion of the elements of the atlantal vertebra,
for a group of Australian taxa (covering 12 genera), but
also stated as including taxa from New Guinea and New
Caledonia (though these were not specifically listed). The
Pseudemoia group as conceived by Greer comprised an
extensive suite of taxa centred on the Coral and western
Tasman Seas, but explicitly excluded part of the New
Caledonian skink fauna, and in doing so argued for non-
monophyly of the group of endemic New Caledonian skink
lineages. Concurrently Hutchinson etal. (1990) reviewed the
Australian species previously assigned to Leiolopisma using
primarily genetic criteria. This study further promoted the
dismembering of the large, still polyphyletic “ Leiolopisma”
as it existed at the time, by transferring the Australian species
formerly in the genus to five (four newly erected) putatively
monophyletic (endemic) genera, and suggesting Leiolopisma
be restricted to the type species telfairii. The genera proposed
by Hutchinson et al. (1990) were identified by micro¬
complement fixation comparisons, and supported by a suite
of putative morphological apomorphies consistent with
these genetic groupings. In the process, genetic comparisons
were made with a very limited suite of extralimital taxa that
included two species from New Caledonia, two from New
Zealand, and the type species for Leiolopisma, Scincus
telfairii Desjardin 1831, from Mauritius.
The results of the genetic study by Hutchinson et al.
(1990) had two implications for New Caledonian skink
systematics. Firstly the New Caledonian and New Zealand
species of Leiolopisma were identified as not being
congeneric with, or particularly close to, the Australian
taxa, and neither was the type species telfairii. Secondly,
while the condition of the elements of the atlantal arch was
invariant for the taxa in most of the genera proposed by
Hutchinson et al., it was variable between the species in one
of the genera proposed. Niveoscincus, with eight species
from Tasmania and adjacent south-east Australia, included
one species with the apomorphic condition of having the
atlantal arches of the first cervical vertebrae fused to the
intercentrum (palfreymani), while the remaining species
in the genus had the plesiomorphic condition in which all
elements are separate, in effect suggesting the possibility
of multiple evolutionary events for this character state,
which had been the sole evidence for monophyly of Greer’s
Pseudemoia group.
210
Records of the Australian Museum (2015) Vol. 67
The actions of Hutchinson et al. (1990) in restricting
Leiolopisma to the type species telfairii prompted the
transfer of the residual New Caledonian “ Leiolopisma” to
Lioscincus (type species Lioscincus steindachneri Bocage,
1873) by Bauer & Sadlier (1993), the only available
generic name among the constituent species. The lack of
apomorphies defining Lioscincus and its probable polyphyly
were explicitly acknowledged at that time, the action being
a nomenclatural necessity rather than an inference of intra¬
generic cohesiveness. For the same reason the New Zealand
“ Leiolopisma ” were transferred to Oligosoma by Patterson
and Daugherty (1995). These actions have resulted in the
concept of a significantly redefined Leiolopisma restricted
to a single extant taxon, the type species telfairii , and the
fossil species mauritianus from the Mascarene Islands
(Arnold, 1980), and the fossil species ceciliae from Reunion
(Arnold & Bour, 2008). However, one species, described as
Leiolopisma alazon Zug in 1985, has not been considered in
these recent studies. The species occurs on the outer islands
of Fiji, and until recently was only known from the type
series, and no tissue samples have been available for genetic
studies. It currently resides within the Lygosominae (Zug,
2013), but otherwise its affinities remain obscure.
Taxonomy
A revised taxonomy for “ Lioscincus ”
Recent investigation of intergeneric relationships among
the members of the Eugongylus group of skinks has been
through the use of mitochondrial and nuclear DNA sequence
data. The scheme of relationships (Fig. 1) resulting from the
study by Smith et al. (2007) strongly supported monophyly
of all the morphologically based New Caledonian genera
previously recognized (Sadlier, 1987; Sadlier & Bauer,
1997; Sadlier et al., 1997, 2004b, 2006b) exclusive of
“ Lioscincus ”, and identified four independent and unrelated
genetic lineages within “ Lioscincus
• Leiolopisma tillieri Ineich & Sadlier, 1991 + Lioscincus
maruia Sadlier, Whitaker & Bauer, 1998.
• Lygosoma (Leiolopisma) novaecaledoniae Parker, 1926.
• Lygosoma (Mocoa) nigrofasciolatus Peters, 1869.
• Lioscincus steindachneri Bocage, 1873 + Lioscincus
vivae Sadlier, Bauer, Whitaker & Smith, 2004.
The lineage “ Lioscincus ” maruia + “ Lioscincus ” tillieri ,
and the species “ Lioscincus ” novaecaledoniae , were each
found to have no obvious affinities to other New Caledonian
skink genera or to each other. Within a well-supported group
that included all other New Caledonian taxa, a sister group
relationship between Lioscincus steindachneri + Lioscincus
vivae (= Lioscincus s.s .) and Celatiscincus was identified,
while “ Lioscincus ” nigrofasciolatus was nested within a
poorly supported group that also included Phoboscincus (as
represented by garnieri ), and Lacertoides.
Genetic analyses presented subsequently by Chappie et
al. (2009), Caut etal. (2013) and Ineich et al. (2014), while
either involving less thorough sampling of New Caledonian
genera, or fewer genes, have similarly found no evidence
for monophyly of any combination of these four Lioscincus
groups.
“ Lioscincus ” novaecaledoniae was found to be the
outgroup to the lineage of all other New Caledonian genera
plus the New Zealand Oligosoma by Ineich et al. (2014).
The position of “Lioscincus ” tillieri + “Lioscincus ” maruia
(Smith etal., 2007; Ineich etal. 2014) has varied in different
studies, but it has never been recovered as sister to other
Lioscincus alone nor tillieri when included without its sister
species (Chappie et al., 2009). The study by Ineich et al.
(2014), using only two of the genes used by Smith et al.
(2007), similarly found this group to be closer to the New
Zealand Oligosoma than to the New Caledonian radiation,
but with low support. In contrast, Chappie et al. (2009), using
five mitochondrial genes and one nuclear gene, but with
just four New Caledonian genera represented, found tillieri
to nest deeply among the other New Caledonian genera,
as sister to Caledoniscincus. Caut et al. (2013), using two
mitochondrial genes, neither used by Smith et al. (2007),
similarly found tillieri to nest among the New Caledonian
genera, but as sister to Phoboscincus + nigrofasciolatus ,
with Caledoniscincus outside this (Caut et al., 2013).
“ Lioscincus” nigrofasciolatus was recovered as sister to
Phoboscincus by Caut et al. (2013), but with low support,
and Lioscincus steindachneri has consistently been recovered
as sister to Celatiscincus (Ineich et al., 2014).
Similarities in aspects of morphology and behaviour
between “ Lioscincus ” tillieri and “ Lioscincus ” maruia , and
by inference a sister species relationship, had previously
been suggested (Sadlier etal., 1998; Bauer & Sadlier, 2000).
However, the suite of morphological apomorphies shared by
these two species was at that time not considered sufficiently
compelling alone to diagnose them as a monophyletic lineage
exclusive of other “ Lioscincus ” taxa.
The distinctiveness of “Lioscincus” novaecaledoniae had
long been recognized, as had the problematic nature of its
relationships with regard to other New Caledonian skinks.
Its blue mouth colour (Sadlier, 1987), a feature not present
in any other New Caledonian skink, was not considered
sufficient by itself to identify the species as a lineage
warranting generic recognition. Subsequent morphological
studies (Sadlier, 2010) have identified a broader suite of
apomorphies that further diagnose the species, and which
in combination with a high level of genetic differentiation
indicate the species warrants recognition as a separate genus
but with no clear indication as to its relationships.
A sister taxon relationship between “ Lioscincus”
nigrofasciolatus and “ Lioscincus ” greeri has also long been
recognized and diagnosed on morphological criteria (Sadlier,
1987; Bauer & Sadlier, 2000). The latter species is known
only from the original specimen(s) collected (Bohme, 1979;
Meier, 1979) and no tissue samples were available for the
study by Smith et al. (2007). Recognition of these taxa as
a lineage warranting generic rank had been tempered by
uncertainty regarding its affinities with a morphologically
similar subset of species in the Pacific region—genus Emoia
(the samoensis group, Greer pers. comm.)—a problem that
no longer exists with the recognition of a monophyletic
New Caledonian skink radiation inclusive of “ Lioscincus ”
nigrofasciolatus but exclusive of the Emoia samoensis group
(as represented by Emoia loyaltiensis , Smith et al., 2007).
The close genetic relationship of the species Lioscincus
steindachneri and Lioscincus vivae indicated by genetic
analysis (Smith et al., 2007) was unexpected. The two species
are dissimilar in their biology and overall appearance, but
share a limited suite of morphological apomorphies (Sadlier
et al., 2004a) that supported the relationship retrieved in the
Sadlier et al .: New Caledonian lizard genus Lioscincus
211
- 1/94
JQt
.65/-
Panaspis africana
- Cryptoblepharus novocaledonicus
- Emoia cyanura
- Menetio greyii
- Eugongylus rufescens
- Err
rm-
Emoia loyaltiensis
-Leiolopisma telfairii
|54C
fh
M-
- Morethia adelaidensis
- Pseudemoio pagenstecheri
- Saproscincus rosei
-Lampropholis guichenoti
■ Niveoscincus pretiosus
- Lygisaurus foliorum
1/75
- Cautulo zia
- 1^Q Q-|~ | Lioscincus novocaledoniae = Caesoris gen. nov.
1/100
.58/-
1 /-
1/93
V-
| Lioscincus maruia
1/100 -f | Lioscincus tlllieri
- Oligosoma lichenigero
-™> 8 'iftzzs'r
= Phasmasaurus gen. nov.
707=
Oligosoma aenea
1 / 100 ,
- Oligosoma suteri
.99/—
.95/—
1/69
t \Nannosdncus garrulus
- Nannoscincus gracilis
(— Nannoscincus gracilis
d-A/
1/96
— 11/68
1 /100 L Nannoscincus sfevini
__t- Nannoscincus mariei
—I- '4/-- Nannoscincus fuscus
•81/- i Ar//>o Nannoscincus greeri
Nannoscincus humectus
X95763”
— .84/-
.99/-
flTTOO
.89/-
.95/-
1 7 oq ivunnusuiiLus numecius
Nannoscincus hanchisteus
— Sigaloseps deplanchei
—Sigaloseps ruficauda
Celatiscincus similis
Celatiscincus euryotis
11/100
1/87
Lioscincus vivae
| Lioscincus steindachneri
= Lioscincus s.s.
.94/-
.94/-
| Lioscincus nigrofasciolatus = Epibatorgen nov.
1 / 1 qq [~ | Lacertoides pardalis
1 /iooF I Phoboscincus gamieri
- mxL
.81/-
[1/73
1/97
.85/-
| Kanakysaurus viviparus
Marmorosphax taom
Marmorosphax boulinda
.1/10 0 | Marmorosphax tricolor
—| Marmorosphaxmontana
- Tropidoscincus aubrianus
1/100 | - Tropidoscincus boreus
1/10041 I Tropidoscincus variabilis
.66/65
{_ | Graciliscincus shonae
V-
1/100
.54/-
- Phaeoscincus taomensis
- Simiscincus aurantiacus
.58/58
1/93
Caledoniscincus orestes
- ll/87 | Caledoniscincus atropunctatus
Caledoniscincus chazeaui
.94/56 1/100 1 — Caledoniscincus terma
_ n-prrchCaledoniscincus auratus
1 - Caledoniscincus renevieri
- Caledoniscincus festivus
1/74 rr ^-.— Caledoniscincus austrocaledonicus
1/71
.94/67
- Caledoniscincus austrocaledonicus
- Caledoniscincus haplorhinus
Figure 1. Phylogenetic relationships of the New Caledonian skinks within the Eugongylus group of skinks as determined by
DNA sequence data (after Smith et al ., 2007) based on combined partitioned Bayesian analysis of one mitochondrial (ND2)
and two nuclear genes (c -mos & Rag-1) with the polyphyly of “ Lioscincus ” highlighted. Support values are Bayesian posterior
probability/bootstrap values, support less than P p 0.50 or bootstrap 50% not shown.
212
Records of the Australian Museum (2015) Vol. 67
molecular study. The placing of Lioscincus steindachneri
+ Lioscincus vivae (= Lioscincus s.s.). by the genetic data
in a well-supported sister relationship with Celatiscincus
is supported by these taxa sharing a morphological
apomorphy, divided nuchal scales, not commonly seen in
other Eugongylus group taxa, and further reinforces the
distinctiveness and relationships of this revised and more
limited Lioscincus from the other taxa formerly included
in Lioscincus.
It is also worth noting here that in the study by Smith et
al. (2007), Phoboscincus was represented by only a single
species, P. garnieri, and monophyly of the genus inclusive of
that species was called into question by Ineich (2009), based
primarily on morphological criteria. Subsequent inclusion
of the recently rediscovered P. bocourti into the Smith et al.
(2007) tree supported monophyly of the genus inclusive of
both taxa (Ineich et al. 2014).
The redefinition and restriction of Lioscincus and
erection of three new genera proposed here to accommodate
the genetic lineages identified by Smith et al. (2007)
for the species nigrofasciolatus, tillieri + maruia , and
novaecaledoniae (see Fig. 2 for representatives of these
lineages) draw substantially from genetic data to support
these each being distinctive evolutionary entities. Although
we are reluctant to erect new genera to accommodate one
or two species, the data at hand suggest this as the most
prudent course of action. As noted above, Lioscincus as
presently conceived, is demonstrably non-monophyletic
and to retain this name for the current constituent species
would be misleading. Affinities of three of the lineages of
“Lioscincus ” are obscure or poorly supported, suggesting
that the initial radiation of the New Caledonian skinks
was rapid and to date neither the addition of more taxa or
more genes has clarified these relationships. To link these
clades with any other genus or with one another through
the use of a shared generic name would be baseless, and
even in the case of Lioscincus s.s. + Celatiscincus , where
affinities are well-supported, deep genetic divergence and
contrasting sets of apomorphies are more consistent with
generic level differences typical in the Eugongylus group.
Diagnoses for these generic arrangements are derived from
an expanded suite of morphological characters (Sadlier,
2010: appendix 1), and are here applied to these newly
recognized lineages identified by the genetic studies. In this
respect the morphological data does not in itself provide an
independent test for monophyly of these genetic lineages,
but rather serves to provide further support for the genetic
lineages being natural groups of species on independent
evolutionary trajectories. Similar approaches to the use
of morphology were made by Hutchinson et al. (1990) in
providing characters to diagnose evolutionary units identified
by genetic studies within the Australian Eugongylus group
skinks, within the Australian skink genus Egernia (Gardner
et al ., 2008) and more recently within the Sphenomorphus
group skink genus Eulamprus (Skinner et al. , 2013), to which
generic names were assigned in each case.
An asterisk indicates the apomorphic character state for
the genus within the context of the Eugongylus group skinks
as determined by Sadlier (2010) in the following generic
diagnoses.
Genus Lioscincus Bocage
Lioscincus Bocage, 1873: 228.
Type species. Lioscincus steindachneri Bocage, 1873: 228.
Diagnosis. The species of Lioscincus s.s. are moderately
large in size (maximum snout vent length [SVL]: L. vivae 55
mm; L. steindachneri 88.5 mm) with a stout body, moderate¬
ly well-developed limbs and digits, and a relatively long tail
(maximum tail length: L. steindachneri 180% SVL; L. vivae
210% SVL). The ear opening is large and lacks obviously
enlarged lobules around the anterior edge.
Scalation (Fig. 3): * distinct supranasal scales absent; nasal
scale lacking a postnasal suture (a crease is sometimes
present in steindachneri ); frontonasal broader than long;
prefrontals large and narrowly to moderately separated;
frontal elongate, longer than broad; supraoculars four;
* frontoparietals fused to form a single scale; interparietal
distinct; * nuchal scales divided so that the parietals are each
bordered by three scales comprising a divided nuchal (two
similar sized scales) and the upper secondary temporal scale;
primary temporal single; lower secondary temporal single;
tertiary temporals two; postlabials two; * anterior loreal
reduced to a semilunar scale positioned on the anterodorsal
margin of the nasal and either failing to, or only narrowly
contacting, the labials; supraciliaries usually seven; upper
labials seven with the fifth subocular and either contacting
the lower eyelid {steindachneri) or separated by a complete
row of subocular scales (vivae), postmental contacting first
and second lower labial; transversely enlarged chinshields
three, first pair in broad contact, second pair separated by
one scale, third pair separated by three scales, all chinshields
contacting the lower labials; dorsal scales of body smooth.
Osteology, premaxillary teeth 11; atlantal arches and
intercentrum of first cervical vertebrae present as three
separate units but showing some degree of partial (vivae) or
superficial (steindachneri) fusion; presacral vertebrae usually
29; phalangeal formula for the manus 2.3.4.5.3 and for the
pes 2.3.4.5.4; two pairs of mesosternal ribs.
The suite of apomorphic character states identified
above will distinguish a redefined Lioscincus from all other
genera in the Eugongylus group of skinks, including the
new genera described here. In particular the combination of
fused frontoparietal scales and division of the nuchal scale
(such that three scales border each of the parietal scales)
serves to distinguish Lioscincus from all genera outside
the endemic New Caledonian skink radiation. Within the
endemic New Caledonian skink radiation only Celatiscincus ,
Kanakysaurus and Phoboscincus bocourti also share this pair
of scalation apomorphies (Sadlier, 2010), although division
of the nuchal scale occurs variably in Simiscincus. The
nuchal scale division in Kanakysaurus is regular but part of
a more extensive pattern of fragmentation of scales in the
temporal and nuchal region, and fragmentation of the nuchal
and temporal scales in Phoboscincus bocourti is extreme and
irregular and also part of a pattern of fragmentation of scales
in the temporal and nuchal region. By contrast division of
the nuchal scale in Lioscincus and Celatiscincus is regular
and not associated with a more extensive pattern of head
shield fragmentation. As such, division of the nuchal scales to
varying degrees has likely evolved independently in several
Sadlier et al .: New Caledonian lizard genus Lioscincus
213
Figure 2. Species now included in Lioscincus s.s., (a) L. steindachneri and (b) L. vivae; and representatives of species included in the
genera: Phasmasaurus, (c) P. tillieri and (d) P. maruia ; Epibator, (e) E. nigrofasciolatus, and Caesoris, (f) Caesoris novaecaledoniae.
of the major groups of endemic New Caledonian skinks,
with its presence in Lioscincus and Celatiscincus a putative
synapomorphy for a sister relationship between these genera.
Celatiscincus has two morphological apomorphies not shared
with Lioscincus , an elevated number of premaxillary teeth
(13 vs 11) and the body scales with weak keels (vs smooth).
Etymology. Bocage (1873) does not give the origin of the
name Lioscincus , but it is presumably derived from the Greek
leios (smooth), and in allusion to the unkeeled scalation of
the type species steindachneri.
Intergeneric relationships. The sister group relationship
between Lioscincus s.s. and Celatiscincus inferred by the
genetic data (Smith et al., 2007) is supported by these
taxa sharing the apomorphic condition for three scalation
characters, the combination of which is unique within the
Eugongylus group of skinks: fusion of the frontoparietal
scales; parietals each bordered by at least three scales, two of
which are a divided nuchal scale and one the upper secondary
temporal; and a narrowing of the anterior loreal basally to
the point where it can be occluded from contact with the
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Records of the Australian Museum (2015) Vol. 67
Figure 3. Head scalation of Lioscincus s.s. as seen in L. steindachneri (adapted from Sadlier, 1987) showing some of the primary
diagnostic features for the genus including fused frontoparietal scales (FP), a narrow-based and semilunar anterior loreal scale
(AL) which contacts the upper labial scales narrowly or not at all, and divided nuchal scales (N), compared to the plesiomorphic
condition of a broad-based anterior loreal and single transversely elongate nuchal either side of the neck as seen in Epibator
(Fig. 4), Phasmasaurus (Fig. 5) and Caesoris novaecaledoniae (Fig. 7), and paired frontoparietals seen in Epibator (Fig. 4).
labials. Celatiscincus is distinguished from Lioscincus s.s.
in having an elevated number of premaxillary teeth (13 vs
11). The relationships of Geoscincus, an unusual monotypic
genus of skink known only from the two type specimens
collected in 1975 (Bohme, 1976), are not available from
any of the genetic analyses and its relationships to other
Eugongylus group skink remain obscure. Regardless,
Geoscincus is readily distinguished from Lioscincus s.s. in
having highly fragmented posterior head shields, including
division of the upper secondary temporal scale, a greatly
reduced number of premaxillary teeth (6-9 vs 11), and in
having the plesiomorphic condition of divided (vs fused)
frontoparietal scales.
Recognized species. Two, Lioscincus steindachneri Bocage
and Lioscincus vivae Sadlier, Bauer, Whitaker & Smith.
Sadlier et al .: New Caledonian lizard genus Lioscincus
215
Lioscincus steindachneri Bocage, 1873: 228
Distribution. Panie Range in the far north-east, and central
metamorphic ranges as far south as Me Adeo.
Comments. Recorded only from humid forest habitat.
Lioscincus vivae Sadlier, Bauer,
Whitaker & Smith, 2004: 211
Distribution. The central-west ultramafic massifs of
Kopeto and Paeoua.
Comments. Recorded from the edge of high elevation
closed forest and throughout maquis shrubland to as
low as 500m elevation (Whitaker, 2006).
Intrageneric relationships. There is a high level of support
for L. steindachneri and L. vivae as sister taxa (BPP 0.95)
in the combined mitochondrial ND2 and nuclear RAG1 +
c -mos molecular phylogeny (Smith et al., 2007), but with
substantial nucleotide sequence divergence between these
taxa for the mitochondrial ND2 gene of around 12.5% (a
similar level of sequence divergence was found between
the two species of the sister-genus Celatiscincus). Ineich et
al. (2014) similarly found a high level of support (BPP 1.0)
for the sister-species relationship. The high level of genetic
differentiation between the two taxa is complemented by the
differences in morphology previously identified, and clearly
supports their recognition as divergent and independent
evolutionary entities. The differences in morphology are
substantial, such that their sister relationship was not
immediately obvious. The two species differ markedly in
overall appearance, most notably in the absence of sexual
dichromatism in L. steindachneri , whereas adult male
L. vivae have a different coloration to adult females and
juveniles of that species. They also differ significantly
in distribution and habitat preferences. Lioscincus
steindachneri is restricted to moist habitat in humid forests
on non-ultramafic soils of the north-east and east-central
ranges, whereas Lioscincus vivae occurs primarily in maquis
shrubland on the west coast ultramafic ranges of Massif de
Kopeto and Paeoua (Whitaker et al ., 2004).
Epibator gen. nov.
Type species. Lygosoma ( Mocoa) nigrofasciolatum Peters,
1869: 435 designated hereby.
Diagnosis. The species are moderately large to large in size
(maximum SVL range 61 mm for greeri and 112 mm for
nigrofasciolatus) with a moderately elongate body, long
limbs and digits, and a moderately long tail (maximum tail
length 145 and 180% of SVL, respectively).
Scalation. *no distinct supranasal; nasal scale with a
prominent postnasal suture or crease; frontonasal as broad
as long; prefrontals large and narrowly separated or in
narrow contact; frontal longer than broad; supraoculars four;
frontoparietals paired; interparietal distinct; parietals each
bordered by a single nuchal and upper secondary temporal
scale; ^primary temporals usually two; lower secondary
temporal single; tertiary temporals two; loreals two, each
contacting the labials broadly; supraciliaries 7 (greeri)
or 8 (i nigrofasciolatus ); * upper labials 8 or more with the
sixth subocular and contacting the lower eyelid, and the
Hast divided by an oblique suture into an upper and lower
scale (Fig. 4); postmental contacting first and second lower
labial; * enlarged chinshields 4-5, first pair in broad contact
medially, second pair in moderate contact, first pair usually in
partial contact with lower labials but * remaining chinshields
separated from lower labials by 1-2 rows of smaller scales
(Fig. 4); body scales smooth and moderately small (34^12
longitudinal rows at midbody); * basal dorsal scales of the
toes divided.
Osteology, premaxillary teeth 11; atlantal arches of first
cervical vertebrae and intercentrum present as three separate
units; 29 presacral vertebrae; a phalangeal formula of
2.3.4.5.3 for the manus and 2.3.4.5.4 for the pes; two pairs
of mesosternal ribs.
The suite of apomorphic character states identified above
will distinguish Epibator gen. nov. from all other genera in
the Eugongylus group of skinks, including the new genera
described here. Within the endemic New Caledonian skink
radiation the presence of 8 or more upper labials with the
last divided by an oblique suture and the separation of the
chinshields from the lower labial scales by 1-2 rows of small
intervening scales will distinguish Epibator gen. nov. from
all other genera except Phoboscincus. Epibator gen. nov.
is distinguished from Phoboscincus in having no distinct
supranasal scale whereas the nasal scale of Phoboscincus
is distinctly divided into three separate elements, the
plesiomorphic condition. Further, Phoboscincus has two
apomorphic character states not present in Epibator gen.
nov., fused (vs paired) frontoparietal scales and enlarged
fang-like (vs peg-like) anterior teeth.
Etymology. From the Greek for climber (epi, upon + bates ,
one that walks or haunts), alluding to the arboreal habits of
the type species. The gender is masculine (Brown, 1956).
Intergeneric relationships. The molecular data presented
by Smith et al. (2007) retrieved a phylogeny which placed
Epibator gen. nov. (as represented by nigrofasciolatus)
within a subgroup containing Epibator gen. nov. as the sister
to Lacertoides + Phoboscincus. This grouping received
a moderate to low level of support (BPP 0.94; bootstrap
values <50%), and the species Phoboscincus bocourti and
Epibator greeri were missing from the analysis. However,
there are no obvious shared apomorphies in morphology
between these genera, though there is a tendency towards
extreme fragmentation of the scales in the temporal region
in all genera. However, this trait is also seen in a number of
other members of the Eugongylus group, particularly larger
taxa e.g. Eugongylus species and Oligosoma grande (Sadlier,
2010), and it is possible its occurrence in the subgroup that
includes Phoboscincus , Epibator gen. nov. and Lacertoides,
could be independently derived.
The genetic grouping of Phoboscincus , Lacertoides , and
Epibator gen. nov., or an inference of relationship between
any of these genera, has not previously been suggested.
Limited support for this grouping comes from the three
genera sharing two scalation characters which could be
considered as putative apomorphies to define the subgroup:
an elevated number of 8 or more upper labial scales; and
division of the basal dorsal toe scales, a derived trait that has
evolved independently in several groups of endemic New
Caledonian skinks (see Sadlier, 2010: appendix 1). Species
in all three genera also share an additional apomorphy in
having two (rather than a single) primary temporal scales,
though this trait is also present in Kanakysaurus and a range
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Records of the Australian Museum (2015) Vol. 67
Figure 4. Head scalation of Epibator as seen in E. nigrofasciolatus (adapted from Sadlier, 1987) showing some of the primary diagnostic
features for the genus including a divided primary temporal scale (PT) and the majority of chinshields (CH) separated from lower labials
by 1-2 rows of smaller scales, compared to the plesiomorphic condition of a single enlarged primary temporal scale and the enlarged
chinshields contacting the lower labials as seen in Epibator (Fig. 4), Phasmasaurus (Fig. 5) and Caesoris novaecaledoniae (Fig. 7).
of other more distant taxa in the Eugongylus group. A more
recent genetic study (Ineich et al., 2014) that included
both Phoboscincus bocourti and Phoboscincus garnieri
also recovered a relationship of E. nigrofasciolatus with
Phoboscincus, although that study used one less nuclear
gene, had lower support values for the relationship of these
two taxa (BPP 0.53, bootstrap values <50%), and Lacertoides
was not identified as part of this lineage. Rather, the genetic
study of Ineich et al. retrieved a relationship for Lacertoides
with Kanakysaurus and Marmorosphax, but with no support
for relationships between these genera and only low support
for the three genera as a subgroup (BPP 0.53, bootstrap
values <50%).
Epibator gen. nov. and Phoboscincus share two
morphological apomorphies, separation of the chinshields
from the lower labial scales by 1-2 rows of small intervening
scales and division of the last upper labial obliquely, that
argue for a sister taxon relationship between these genera,
contrary to the genetic evidence of Smith et al. (2007) which
retrieves Lacertoides as the sister to Phoboscincus, but in
accordance with the genetic evidence of Ineich etal. (2014).
Separation of the chinshields from the lower labials by an
intervening row of scales is variable within Phoboscincus,
with the chinshields of bocourti completely separated
from the lower labials, but only partially (third chinshield
only) in garnieri. These traits have only otherwise been
Sadlier et al .: New Caledonian lizard genus Lioscincus
217
recorded from the monotypic Geoscincus (Sadlier, 1987),
a genus whose phylogenetic affinities are obscure and for
which genetic data are unavailable. Geoscincus is readily
distinguished from Epibator gen. nov. in having a highly
reduced number of premaxillary teeth (6-9 vs 11), and a
“scaled” lower eyelid, the homology of which is unclear and
is otherwise only seen in the New Caledonian skink radiation
in Phoboscincus boconrti. By comparison, Lacertoides has
the plesiomorphic condition of all three chinshields fully in
contact with the lower labials and an undivided last upper
labial.
Recognized species. Two, Epibator nigrofasciolatus (Peters)
and Epibator greeri (Bohme).
Epibator nigrofasciolatus (Peters)
Synonyms. Lygosoma (Mocoa) nigrofasciolatum Peters,
1869: 435. Lygosoma arborum Bavay, 1869: 19.
Lygosoma deplanchei Bocage, 1873: 229 (non
Lygosoma deplanchei Bavay, 1869 = Sigaloseps
deplanchei).
Distribution. Widespread throughout New Caledonia,
including large and small offshore islands, and the
Loyalty Islands.
Comments. Recorded from a wide range of habitats at all
elevations.
Epibator greeri (Bohme)
Synonym. Leiolopisma greeri Bohme, 1979: 140.
Distribution. Koumac region in the north-west of Grande
Terre.
Comments. Bohme described the species from a single
individual collected in forest. It has not been seen
with certainty since, although live individuals similar
in colouration to the type of Epibator greeri have
been photographed (by one of us, RS) from southern
Grande Terre.
Genus Phasmasaurus gen. nov.
Type species. Leiolopisma tillieri Ineich & Sadlier, 1991
designated hereby.
Diagnosis. Moderately large in size, maximum SVL 61
mm ( maruia ) and 64 mm ( tillieri ), with a moderately
elongate body, well developed limbs and digits, and very
long tail* (maximum tail length range c. 250-300% of SVL
respectively).
Scalation (Fig. 5): distinct supranasal absent*; nasal scale
lacking a prominent postnasal suture; frontonasal broader
than long; prefrontals large; frontal longer than broad;
supraoculars four; *frontoparietals fused; interparietal
distinct; parietals each bordered by a single nuchal and one
{maruia) or two {tillieri) upper secondary temporal scales;
primary temporal single; lower secondary temporal single;
tertiary temporals usually two; nasals widely separated;
supraciliaries usually seven {maruia) or fewer {tillieri),
reduction through fusion; upper labials usually seven with the
fifth subocular and contacting the lower eyelid; postmental
contacting first and second lower labial; chinshields three,
first pair in broad contact, all in contact laterally with the
lower labial scales; *body scales keeled {maruia three weak
keels; tillieri two strong keels).
Osteology, premaxillary teeth 9 {tillieri) or 11 {maruia)',
atlantal arches and intercentrum of first cervical vertebrae
present as three separate units; 29 presacral vertebrae;
*postsacral vertebrae c. 60+; phalangeal formula for the
manus of 2.3.4.5.3 and for the pes of 2.3.4.5.4; two pairs of
mesosternal ribs.
Reproduction mode', variable within the genus with one
species oviparous (egg laying: maruia) and one viviparous
(live-bearing: tillieri).
The suite of apomorphic character states identified
above is modest, but will distinguish Phasmasaurus gen.
nov. from all other genera in the Eugongylus group of
skinlcs, including the new genera described here. The key
morphological apomorphy uniting tillieri and maruia as
sister taxa, and which serves to distinguish them from
as a distinct evolutionary entity from most Eugongylus
group genera is the exceptionally long tail of the species of
Phasmasaurus, characterized by a high postsacral vertebrae
number of 60 or more. This character state is otherwise only
seen within the Eugongylus group in some Emoia (species
in the samoensis group), which are outside the endemic
New Caledonian skink radiation. Phasmasaurus gen. nov.
can be distinguished from these Emoia in having fused {vs
paired) frontoparietals, keeled {vs smooth) body scales, and
fused (vs- distinct) supranasal scales, although the evolution
of supranasal scales within the Eugongylus group, and the
polarity of their presence are open to question (Sadlier, 2010).
Etymology. From the Greek Phasma, a spectre, in the
sense of the insect genus Phasma Lichtenstein, type genus
for the family Phasmatidae, and for the order Phasmatodea
in general, alluding to the phasmid-like appearance and
behaviour of species in the group. Gender of the generic
name based on the Greek sauros (masculine).
Intergeneric relationships. Support for the two species
included in Phasmasaurus as a genetic lineage in the study
by Smith et al. (2007) based on one mitochondrial (ND2) and
two nuclear (c -mos & RAG-1) genes was high (BPP1 but with
Bootstrap support <50%), and a similar, but slightly lower
level of support (BPP 0.95%, bootstrap 71%) was recovered
by Ineich et al. (2014). The two species in Phasmasaurus
share only a modest suite of morphological apomorphies
that serve to define it, including fused frontoparietal scales,
keeled body scales, a windowed lower eyelid and a very long
tail, all character states that can be found in some other taxa
in the Eugongylus group. Additional support for monophyly
of the two species as a lineage (outside of the genetic data)
comes from unusual shared attributes in habitat preference
and behaviour. The strict habitat preference of both species
to maquis shrubland and shared unusual aspects of behaviour
(Sadlier et al., 1998) in combination, lend further support to
the genetic data for these taxa comprising an independent
evolutionary lineage highly divergent from others in the
Eugongylus group of skinks.
The scheme of relationships for the Eugongylus group
skinks presented by Smith et al. (2007) placed the species
here included in Phasmasaurus (and the New Caledonian
species here recognized as Caesoris novaecaledoniae),
outside the well-supported lineage that included nearly
all other endemic New Caledonian skinks, and within a
cluster that included the New Zealand genera inclusive
of the Norfolk/Lord Howe Island Oligosoma lichenigera.
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Records of the Australian Museum (2015) Vol. 67
Figure 5. Head scalation of Phasmasaurus as seen in P. tillieri showing the relatively plesiomorphic configuration of the
majority of head scales except for the fused frontoparietal scales (FP). Note the divided upper secondary temporal (LIST)
an apomorphy for P. tillieri not shared with P. maruia.
although this latter group received no support as a lineage
in its own right. A more recent molecular phylogeny of
Chappie et al. (2009) which sampled extensively the New
Zealand Eugongylus group skinks, and which also included
representative taxa from the New Caledonian skink fauna
for out-group comparison, retrieved a monophyletic
New Caledonian skink group represented by the taxa
Caledoniscincus austrocaledonicus, Nannoscincus mariei ,
Marmorosphctx tricolor , and “ Lioscincus ” tillieri that was
the sister group to the New Zealand + Lord Howe/Norfolk
Island O. lichenigera , and with tillieri now nested within
the (albeit reduced) New Caledonian group. The most
recent molecular phylogeny of Ineich et al. (2009), which
concentrated primarily on the endemic New Caledonian
skink radiation and the genetic relationships of the recently
discovered Phoboscincus bocourti , but which also included
a range of other Eugonglus group genera in the outgroup,
placed the species here included in Phasmasaurus as the
sister to all the endemic New Zealand skinks (also including
Oligosoma lichenigera ), but with no support.
Recognized species. Two, Phasmasaurus tillieri (Ineich
& Sadlier) and Phasmasaurus maruia (Sadlier, Whitaker
& Bauer).
Sadlier et al. : New Caledonian lizard genus Lioscincus
219
Phasmasaurus tillieri (Ineich & Sadlier)
Synonym. Leiolopisma tillieri Ineich & Sadlier, 1991: 344.
Distribution. Restricted to the southern ultramafic ranges
of New Caledonia.
Comments. Endemic to maquis shrubland habitats.
Phasmasaurus maruia (Sadlier, Whitaker & Bauer)
Synonym. Lioscincus maruia Sadlier, Whitaker & Bauer,
1998; 335.
Distribution. East-central region ultramafic ranges and the
west-central ultramafic ranges as far north as Plateau
de Tia and Massif de Kopeto.
Comments. Restricted to maquis shrubland habitats.
Intrageneric relationships. Two genetic-based studies
(Smith et al., 2007; Ineich et al., 2014) retrieve P.
tillieri and P. maruia as highly supported (BPP 1.0)
sister taxa. However, there is also substantial genetic
and morphological differentiation between the two taxa.
Nucleotide sequence divergence between P. tillieri and P.
maruia for the mitochondrial gene ND2 was around 20.0%,
approaching the level of genetic distance found between
some genera of New Caledonian skinks (Sadlier & Bauer,
unpublished). The high level of genetic differentiation
between the two taxa is complemented by a number of
differences in morphology and clearly supports their
recognition as independent evolutionary entities. While
the two species are similar in overall body form in both
having long legs, a long tail and an angular head, there
are substantial differences in scalation. In particular, the
degree of morphological differentiation seen in tillieri , as
expressed in the number of species-specific apomorphies,
is exceptional in comparison to most other New Caledonian
skinks. In particular the strongly bicarinate scales with the
keel interrupting the posterior free edge is a trait unique to
the species within the context of the Tasmantis group skinks
(Fig. 6a). By comparison maruia is plesiomorphic in its
head scalation and the weakly tricarinate body scales (Fig.
6b) of this species likely represent the more plesiomorphic
form of this character for the genus (see Sadlier, 2010:
appendix 1). The two species also differ markedly in
reproductive mode, with P. tillieri giving birth to live young
while P. maruia has retained the plesiomorphic egg-laying
condition. This is the only genus in the Eugongylus group
where both modes of reproduction occur. The extent of
genetic and morphological differentiation between P.
tillieri and P. maruia raises the question of recognising
each as a distinct monophyletic genus. While a convincing
stand-alone diagnosis could be made for tillieri , the
morphological diagnosis for maruia as a lone entity would
be weak and lacking in unique apomorphies to further
differentiate it from tillieri. For these reasons this action
has not been proposed at this time and their inclusion as
sister taxa under Phasmasaurus rests primarily on the
implications of the phylogenetic relationships retrieved so
far from molecular studies.
In overall ecology the two species share traits in
behaviour and a strict habitat preference only rarely seen
in other endemic New Caledonian skink genera (Sadlier et
al., 1998). Both are restricted solely to maquis shrubland
(.Lacertoides is the only other taxon of skink reliant on this
habitat type), with tillieri widespread across the southern
ultramafic region and maruia on the adjacent central-west
ultramafic ranges. As such, the two species are broadly
parapatric, and they act as ecological analogues within
their respective regions.
Figure 6. SEM image of markedly bicarinate body scales seen in (a) Phasmasaurus tillieri compared to weakly tri-quadricarinate body scales
seen in (b) Phasmasaurus maruia , and compared to the tricarinate body scales seen in (c) Caesoris novaecaledoniae, the plesiomorphic
condition of smooth body scales seen in (d) Epibator nigrofasciolatus and in (e) Lioscincus steindachneri, and the multicarinate body
scales seen in (f) Lioscincus vivae.
220
Records of the Australian Museum (2015) Vol. 67
Figure 7. Head scalation for Caesoris novaecaledoniae (adapted from Sadlier, 1987) showing the relatively plesiomorphic
configuration of the majority of head scales except for the fused frontoparietal scales (FP) and single enlarged tertiary
temporal scale (77), compared to the plesiomorphic condition of paired frontoparietals seen in Epibator (Fig. 4) and two
tertiary temporal scales seen in Epibator (see figure 4), Phasmasaurus (see figure 5) and Caesoris novaecaledoniae (Fig. 6).
Genus Caesoris gen. nov.
Type species. Lygosoma (Leiolopisma) novaecaledoniae
Parker, 1926, designated hereby.
Diagnosis. Maximum SVL 68 mm with a moderately
elongate body, well developed limbs and digits, and
relatively long tail (maximum tail length c. 200% of SVL).
Scalation (Fig. 7): *no distinct supranasal; nasal scale
lacking a prominent postnasal suture; frontonasal slightly
broader than long; prefrontals large; frontal longer than
broad; supraoculars four; * frontoparietals fused; interparietal
distinct; parietals each bordered by a single nuchal and upper
secondary temporal scale; primary temporal single; lower
secondary temporal single; *tertiary temporals fused to form
a single scale; nasals widely separated; supraciliaries usually
seven; upper labials usually seven with the fifth subocular
and contacting the lower eyelid; postmental contacting first
and second lower labial; chinshields three, first pair in broad
contact, all in contact laterally with the lower labial scales;
*body scales with three moderately strong keels dorsally.
Osteology , premaxillary teeth 11; * atlantal arches of first
cervical vertebrae fused to intercentrum; 29 presacral
vertebrae; phalangeal formula for the manus of 2.3.4.5.3
and for the pes of 2.3.4.5.4; two pairs of mesostemal ribs.
Sadlier et al. : New Caledonian lizard genus Lioscincus
221
Reproductive mode, egg laying.
Mouth lining and tongue colour. *deep blue.
The suite of apomorphic character states identified above
will distinguish Caesoris gen. nov. from all other genera in
the Eugongylus group of skinks, including the new genera
described here. In particular, the exceptionally bright blue
mouth colour of Caesoris is unique within the endemic New
Caledonian skink radiation. The presence of a single large
tertiary temporal scale will also serve to distinguish Caesoris
from most other New Caledonian skink genera which have
two tertiary temporal scales, except for Phasmasaurus (a
single enlarged tertiary temporal scale occurs in most P.
maruia) and Lioscincus (a single enlarged tertiary temporal
scale occurs in c. 50% of L. vivae). The species also has an
unusual toe morphology in the context of the Eugongylus
group of skinks in which the basal lamellae of each toe are
broadened laterally compared to those on the distal part of
the toe (Fig. 8), resembling the condition seen in some of
the arboreal geckos in the genus Cyrtodactylus.
Etymology. Derived from the Latin for blue (caesius) and
mouth (oris), in reference to the unique blue mouth colour
of the type species. Gender of name based on “oris”, neuter
(see Brown, 1956).
Intergeneric relationships. The study by Smith et al.
(2007) was unable to identify any close relationship of the
single species in Caesoris to any other genus within the
Tasmantis group of skinks. In this scheme of relationships
the Caesoris lineage lies outside a well-supported clade
of most other endemic New Caledonian skink genera
(Nannoscincus (Lioscincus group + Phoboscincus group
+ Caledoniscincus group), as does the New Caledonian
Phasmasaurus lineage and the lineage that includes the New
Zealand and Norfolk/Lord Howe Island genus Oligosoma.
The most recent molecular phylogeny of Ineich et al. (2014)
placed Caesoris as the sister to all other taxa in the Tasmantis
Clade (= the endemic New Caledonian and New Zealand
skinks combined), but with no support for this placement.
Among the morphological apomorphies that diagnose
Caesoris , a single broad tertiary temporal scale and blue
mouth lining and tongue are unique to the genus within the
context of the taxa included in the endemic New Caledonian
skink fauna. These two characters, in combination with a
broader suite of apomorphies, serve to diagnose Caesoris
from all other genera of skinks in the Tasmantis group, and
support its recognition as a distinct evolutionary lineage as
indicated by the genetic data.
Recognized species. One, Caesoris novaecaledoniae (Parker).
Caesoris novaecaledoniae (Parker)
Synonym. Lygosoma (Leiolopisma) novae-caledoniae
Parker, 1926: 493.
Distribution. Widespread (albeit sparsely) across the
central and northern regions of New Caledonia, with
photographs of live individuals indicating it also
occurs in southern Grande Terre.
Comments. Recorded from a wide range of habitats
including low-mid elevation humid forest and coastal
scrub (Sadlier, 1987), and Acacia scrubland bordering
mesophyll dry forest (Whitaker et al., 2005).
Figure 8. Underside of the (a) manus and (b) pes of Caesoris
novaecaledoniae (AMS R.90454 SVL 61 mm) showing the broad
basal lamellae of all digits and markedly narrower terminal on
the distal-most part of each digit, extending to approximately
the junction of the 3rd and 4th plalange on the 4th toe of the pes,
compared to the lamellae of the (c) manus and (d) pes of Epibator
nigrofasciolatus (AMS R. 177492 SVL 64 mm), also an arboreal
species, which are only moderately broad basally and only extend
to approximately the junction of the 1 st and 2nd plalange on the
4th toe of the pes.
Generic diversity in the endemic New
Caledonian skink fauna
The Eugongylus group of skinks has a primarily Australian/
Pacific region distribution, but with taxa from the sub-Saharan
African region also included (Greer, 1974; Schmitz et al.,
2005; Pyron et al., 2013). The group is currently undergoing
an extensive molecular-based survey of relationships (Shea
et al, unpublished data). It currently stands at 41 recognized
genera, although some (Emoia) are clearly composite, while
the affinities of others (Geoscincus, Tachygyia ) are still unclear
or under debate (Ineich et al, 2014).
The description of the three new genera presented here
brings the number of endemic skink genera present in New
Caledonia to 17 (Table 1). This represents approximately
40% of the generic diversity within the Australian/Pacific
region Eugongylus group of skinks as currently recognised,
and is equivalent to the number of genera in the group
endemic to Australia (c. 14) and New Zealand (1) combined.
The number of endemic Australian genera in the
Eugongylus group of skinks is likely to undergo further
refinement, and their content likely to change, but not
substantially. Regardless, the paradigm of the exceedingly
high level of generic diversity for the endemic New
Caledonian skink fauna when adjusted for area (19,103
km 2 ) compared to other islands in the Pacific region remains
exceptional, particularly when compared to New Zealand
(268,704 km 2 ) which is 14 times its size.
The extent of generic diversity in the endemic New
Caledonian skink fauna reflects the extent of morphological
and behavioural diversity found, which in turn is a reflection
222
Records of the Australian Museum (2015) Vol. 67
Table 1 . The currently recognized endemic New Caledonian
skink genera in order of description.
Year Genus
Author
1872 Nannoscincus
Gunther
1873 Lioscincus
Bocage
1873 Tropidoscincus
Bocage
1974 Phoboscincus
Greer
1987 Geoscincus
Sadlier
1987 Gracilis cine us
Sadlier
1987 Marmorosphax
Sadlier
1987 Caledoniscincus
Sadlier
1987 Sigaloseps
Sadlier
1997 Lacertoides
Sadlier, Shea & Bauer
1997 Simiscincus
Sadlier & Bauer
2004 Kanakysaurus
Sadlier, Smith, Bauer & Whitaker
2006 Celatiscincus
Sadlier, Smith & Bauer
2014 Phaeoscincus
Sadlier, Smith & Bauer
2015 Epibator gen. nov.
Sadlier, Bauer, Shea & Smith
2015 Phasmasaurus gen.
nov. Sadlier, Bauer, Shea & Smith
2015 Caesoris gen. nov.
Sadlier, Bauer, Shea & Smith
of the extent of niche diversity on the New Caledonian
terrain, the end product of which is expressed as an array of
highly divergent lineages, each characterized by a unique
suite of adaptive characteristics.
The initial reallocation of New Caledonian skink taxa
by Sadlier in 1987 to putatively monophyletic genera
resulted in the recognition an additional four genera
(Caledoniscincus, Marmorosphax and Sigaloseps, and a
resurrected Tropidoscincus Bocage, 1873) to accommodate
most of the New Caledonian species formerly assigned to
Leiolopisma. These genera were each defined by a suite of
apomorphic morphological character states, the combination
of which was unique within the Eugongylus group of
skinks. Similarly the monotypic genera Geoscincus (Sadlier,
1987), Graciliscincus (Sadlier, 1987), Lacertoides (Sadlier
et al., 1997), Simiscincus (Sadlier & Bauer, 1997) and
Kanakysaurus (Sadlier et al ., 2004) were recognized and
defined on morphological criteria (although Kanakysaurus
had supporting genetic data). The single species included
in each genus had an array of unusual and highly derived
morphological character states, only a few of which
were shared with taxa in existing genera. Similarly, the
reassessment of Nannoscincus as a strictly endemic New
Caledonian taxon (Sadlier et al ., 2002, 2006), rather than
one shared with Australia, was based on morphological
criteria and monophyly of the genus later supported by
genetic data (Smith et al., 2007). Recognition of “LygosomcT
euryotis as warranting its own genus (Bauer & Sadlier,
2000) was more complex. Although initially recognized as
unique on a combination of morphological and biological
criteria, the erection of Celatiscincus (Sadlier et al ., 2006)
for the species euryotis (and its newly described sister
taxon C. similis) ultimately drew heavily on genetic data
for recognition as a distinct evolutionary entity. By contrast,
the recently proposed Phaeoscincus Sadlier, Smith & Bauer
was initially recognized as a highly divergent evolutionary
lineage on genetic criteria (Sadlier et al ., 2014b). The type
species for the genus ( P. taomensis ) is known from a single
specimen and is markedly similar in morphology to taxa in
Caledoniscincus to which it is the sister, but from which it
is diagnosed by a limited suite of apomorphic characters.
More recently the rediscovery of the enigmatic giant
New Caledonian skink Eumeces bocourti Brocchi, the type
species for the genus Phoboscincus Greer, has permitted a
genetic analysis of the relationships between this species
and its putative congener P. garnieri (Bavay). Phoboscincus
was erected on morphological criteria, and the genetic study
by Ineich et al. (2014) confirms the relationship between
bocourti and garnieri exclusive of all other New Caledonian
taxa, but could not explore a relationship between bocourti
and the giant Tongan species Tachygyia microlepis suggested
earlier by Greer (1974) and later Ineich (2009), as no recent
tissue samples for microlepis were available for analysis.
However, if such a close relationship exists, it would mean
that Tachygyia is also a member of the New Caledonian
skink radiation, implying overwater colonization from New
Caledonia to Tonga.
In summary, most of the endemic skink genera exclusive
of those described here were originally recognized and
defined primarily on morphological criteria, and the
monophyly of these genera (Smith et al ., 2007) and the
content of their constituent taxa have since been supported
by an array of genetic studies (Sadlier et al ., 2009, 2014a).
The initial recognition of these genera on attributes of
morphology, the validity of which has been independently
verified by genetic studies, is testimony to the breadth of
diversity in form and structure within the endemic New
Caledonian skink radiation.
The historical retention of the species nigrofasciolatus
Peters, greeri Bohme, steindachneri Bocage; and novaecal-
edoniae Parker within a “cosmopolitan” Leiolopisma (Sadlier,
1987), assignment of tillieri Ineich & Sadlier (Ineich & Sadlier,
1991) to that genus, and the subsequent assignment of these
taxa to Lioscincus Bocage (Bauer & Sadlier, 1993), reflected
more the conundrum of not being able to assign these taxa
to existing genera without compromising (and weakening)
the integrity of the generic diagnoses of established taxa.
In the case of steindachneri, novae cal edoniae and tillieri
this situation was exacerbated in the past by a lack of
confidence in the (sometimes limited) individual suite
of unique character states (autapomorphies) for each
taxon as sufficient to diagnose them as distinct lineages
on morphological criteria alone. The discovery of new
(sister) taxa for tillieri with the description of Lioscincus
maruia Sadlier, Whitaker & Bauer (Sadlier et al., 1998)
and more recently for steindachneri with the description of
Lioscincus vivae Sadlier, Bauer, Whitaker & Smith (Sadlier
et al., 2004a) provided evidence that each of these species
was now part of a putative multi-species lineage. As such,
the discovery of new taxa related to the species included
in “Lioscincus” , in combination with the acquisition of
extensive genetic data over the past 15 years in particular,
have considerably advanced our understanding of the
affinities of the these species to each other, and to other
New Caledonian skinks, and in doing so realized the true
and extraordinary extent of generic diversity within the
endemic New Caledonian skink fauna.
Sadlier et al. : New Caledonian lizard genus Lioscincus
223
Acknowledgments. Credits for the images used in the figures
presented are as follows: Fig. 1 adapted from Smith et al. 2007;
Fig. 2a,c,d,e photographic images by Ross Sadlier, Fig. 2b by A.
H. Whitaker, and Fig. 2f by Stephane Astrongatt; Figs 3, 4 and
7 drawings are adapted from those produced for publication in
Sadlier, 1987; Fig. 5 originally drawn by Hannah Finley; Fig. 6
scanning electron microscope images and photographic images
for Fig. 8 were produced by Sue Lindsay (Microscopy and
Microanalysis Unit, Australian Museum). The initial genetic studies
that highlighted the polyphyly of Lioscincus were supported by
Grants DEB 0108108 andDEB 0515909 from the National Science
Foundation to A. M. Bauer and T. Jackman.
References
Arnold, E. N. 1980. Recently extinct reptile populations from Maurit¬
ius and Reunion, Indian Ocean. Journal of Zoology 191: 33—47.
http://dx.doi.org/10.1111/j.1469-7998.1980.tb01447.x
Arnold, E. N., and R. Bour. 2008. A new Nactus gecko
(Gekkonidae) and a new Leiolopisma skink (Scincidae) from
La Reunion, Indian Ocean, based on fossil remains and ancient
DNA sequence. Zootaxa 1705: 40-50.
Andersson, L. G. 1908. Two new lizards ( Eurydactylus and
Lygosoma ) from New Caledonia. Arkivfor Zoologi 4(14): 1-5.
Austin, J. J., and E. N. Arnold. 2006. Using ancient and recent DNA
to explore relationships of extinct and endangered Leiolopisma
skinks (Reptilia: Scincidae) in the Mascarene Islands. Molecular
Phylogenetics and Evolution 39: 503-511.
http://dx.doi.org/10.1016/j.ympev.2005.12.011
Bauer, A. M. 1989. Reptiles and the biogeographic interpretation
of New Caledonia. Tuatara 30: 39-50.
Bauer, A. M., and R. A. Sadlier. 1993. Systematics, biogeography
and conservation of the lizards of New Caledonia. Biodiversity
Letters 1: 107-122.
http://dx.doi.org/10.2307/2999755
Bauer A. M., and R. A. Sadlier. 2000. The Herpetofauna of
New Caledonia. Ithaca, New York: Society for the Study of
Amphibians and Reptiles. 310 pp, 24 pis.
Bavay, A. 1869. Catalogue des reptiles de la Nouvelle-Caledonie
et description d’especes nouvelles. Memoires de la Societe
Linneenne de Normandie 15: 1-37.
Bocage, J. V. Barboza du. 1873. Sur quelques sauriens nouveaux
de la Nouvelle Caledonie et de FAustralie. Jornal de Sciencias
Mathematicus Physicas e Naturaes. Lisboa 4: 228-232.
Bohme, W. 1976. Uber die Gattung Eugongylus Fitzinger, mit
Beschreibung einer neuer Art (Reptilia: Scincidae). Bonner
Zoologische Beitrage 27: 245-251.
Bohme, W. 1979. Eine neue Art der Gattung Leiolopisma Dumeril
etBibron, 1839 ausNeukaledonien (Reptilia: Sauria: Scincidae).
Salamandra 15: 140-145.
Brandley, M. C., A. Schmitz, andT. W. Reeder. 2005. Partitioned
Bayesian analyses, partition choice, and the phylogenetic
relationships of scincid lizards. Systematic Biology 54: 373-390.
http://dx.doi. org/10.1080/10635150590946808
Brocchi, M. 1876. Sur un scincoidien nouveau appartenant au
genre Eumeces. Bulletins de la Societe Philomathique de Paris
13(6): 95-97.
Caut, S., M. Holden, M. J. lowers, R. Boistel, and I. Ineich. 2013.
Is Bocourt’s Terrific Skink really so terrific? Trophic myth and
reality. PLoS One 8(10): e78638.
http://dx.doi.org/10.1371/journal.pone.0078638
Chappie, D. G., P. A. Ritchie, and C. H. Daugherty. 2009. Origin,
diversification and systematics of the New Zealand Skink fauna
(Reptilia: Scincidae). Molecular Phylogenetics and Evolution
52: 470-487.
http://dx.doi.org/10.1016/j.ympev.2009.03.021
Desjardins, J. 1831. Sur trois especes de lezard du genre Scinque
qui habitant File Maurice (lie de France). Annales des Sciences
naturelles 22: 292-299.
Gardner, M. G., A. F. Hugall, S. C. Donnellan, M. N. Hutchinson,
and R. Foster. 2008. Molecular systematics of social skinks:
phylogeny and taxonomy of the Egernia group (Reptilia: Scinc¬
idae). Zoological Journal of the Linnean Society 154: 781-794.
http://dx.doi.org/10.1111/j.l 096-3642.2008.00422.X
Greer, A. E. 1970. A subfamilial classification of scincid lizards.
Bulletin of the Museum of Comparative Zoology 139(3): 151-184.
Greer, A. E. 1974. The generic relationships of the scincid lizard
genus Leiolopisma and its relatives. Australian Journal of
Zoology , Supplementary Series 31: 1-67.
http://dx.doi.org/10.1071 /AJZS031
Greer, A. E. 1979. A phylogenetic subdivision of Australian skinks.
Records of the Australian Museum 32(8): 339-371.
http://dx.doi.org/10.3853/j.0067-1975.32.1979.459
Greer, A. E. 1989. The Biology and Evolution ofAustralian Lizards.
Chipping Norton, NSW: Surrey Beatty and Sons Pty. Ltd. 264 pp.
Gunther, A. 1872. On some new species of reptiles and fishes
collected by J. Brenchley, Esq. Annals and Magazine of Natural
History (4)10: 418^120.
http://dx.doi.org/10.1080/00222937208696731
Honda, M., H. Ota, M. Kobayashi, J. Nabhitabhata, H.-S.Yong, and
T. Hikida. 2000. Phylogenetic relationships, character evolution,
and biogeography of the subfamily Lygosominae (Reptilia:
Scincidae) inferred from mitochondrial DNA sequences.
Molecular Phylogenetics and Evolution 15: 452-461.
http://dx.doi.org/10.1006/mpev.l 999.0767
Honda, M., H. Ota, G. Kohler, I. Ineich, L. Chirio, S.-L. Chen, and
T. Hikida. 2003. Phylogeny of the lizard subfamily Lygosominae
(Reptilia: Scincidae), with special reference to the origin of the
New World taxa. Genes and Genetic Systems 78: 71-80.
http://dx.doi.org/10.1266/ggs.78.71
Hutchinson, M. N., S. C. Donnellan, P. R. Baverstock, M. Krieg, S.
Simms, and S. Burgin. 1990. Immunological relationships and
generic revision of the Australian lizards assigned to the genus
Leiolopisma (Scincidae: Lygosominae). Australian Journal of
Zoology 38: 535-554.
http://dx.doi.org/10.1071/Z09900535
Ineich, I. 2009. Bocourt’s terrific skink, Phoboscincus bocourti
Brocchi, 1876 (Squamata, Scincidae, Lygosominae). In Zoologia
Neocaledonica 7, ed. P. Grandcolas. Memoires du Museum
national d’Histoire naturelle 198: 149-174.
Ineich, I., and R. A. Sadlier. 1990. Anew species of scincid lizard
from New Caledonia (Reptilia Lacertilia Scincidae). In Zoologia
Neocaledonica 2, ed. J. Chazeau and S. Tillier. Memoires du
Museum national d’Histoire naturelle 149: 343-347.
Ineich, I., R. A. Sadlier, A. M. Bauer, T. R. Jackman, and S. A.
Smith. 2014. Bocourt’s terrific skink, Phoboscincus bocourti
(Brocchi, 1876), and the monophyly of the genus Phoboscincus
Greer, 1974. In Zoologia Neocaledonica 8, ed. E. Guilbert, T.
Robillard, H. Jourdan, and P. Grandcolas. Memoires du Museum
national d’Histoire naturelle 206: 69-78.
Loveridge, A. 1941. An undescribed skink ( Lygosoma ) from New
Caledonia. Proceedings ofthe Biological Society of Washington
54: 193-194.
Meier, H. 1979. Herpetologische Beobachtungen auf Neukale-
donien. Salamandra 15: 113-139.
Mertens, R. 1928. Neue Inselrassen von Cryptoblepharus boutonii
(Desjardin). Zoologischer Anzeiger 78: 82-89.
Mittleman, M. B. 1952. A generic synopsis of the lizards of the sub¬
family Lygosominae. Smithsonian Miscellaneous Collections
117(17): 1-35.
Parker, H. W. 1926. A new lizard from New Caledonia. Annals and
Magazine of Natural History 18(9): 493-495.
http://dx.doi.org/10.1080/00222932608633546
Patterson, G. B., and C. H. Daugherty. 1995. Reinstatement of the
genus Oligosoma (Reptilia: Lacertilia: Scincidae). Journal of
the Royal Society of New Zealand 25: 327-331.
http://dx.doi.org/10.1080/03014223.1995.9517493
Peters, W. 1869. Uber neue Gattungen und neue oder weniger
bekannte Arten von Amphibien. Monatsberichte Koeniglich
Preussiche Akademie Der Wissenschaften Berlin 1869: 432-445.
224
Records of the Australian Museum (2015) Vol. 67
Peters, W. 1879. Uber Sauroscincus, eine neue Gattung von
Scincoiden, aus Neu-Caledonien. Sitzungs-Bertrichte der
Gesellschaft Naturforschender Freunde zu Berlin 10:149-150.
Pyron, R. A., F. T. Burbrink, and J. J. Wiens. 2013. Aphylogeny and
revised classification of Squamata, including 4161 species of
lizards and snakes. BioMed Central Evolutionary Biology 13:93.
http://dx.doi.org/10.1186/1471 -2148-13-93
Rabosky, D. L., S. C. Donnellan, A. L. Talaba, and I. J. Lovette.
2007. Exceptional among-lineage variation in diversification
rates during the radiation of Australia’s most diverse vertebrate
clade. Proceedings of the Royal Society B 274: 2915-2923.
http://dx.doi.org/10.1098/rspb.2007.0924
Roux, J. 1913. Les reptiles de la Nouvelle-Caledonie et des lies
Loyalty. In Nova Caledonia, Zoologie, ed. F. Sarasin, and J.
Roux, vol. 1(2), pp. 79-160. Wiesbaden: C. W. Kreidels Verlag.
Sadlier, R. A. 1987. A review of the scincid lizards of New
Caledonia. Records of the Australian Museum 39(1): 1-66
(marked 1986 in error on p. 1, published 5 May 1987).
http://dx.doi.org/10.3853/j. 0067-1975.39.1987.164
Sadlier, R. A. 1990. The scincid lizard genus Nannoscincus Gunther:
a revaluation. Memoirs ofthe Queensland Museum 29:487^494.
Sadlier, R. A. 2010. Systematic Studies ofthe Scincid Lizards of New
Caledonia. PhD Thesis, Griffith University, Queensland. 199 pp.
Sadlier, R. A., and A. M. Bauer. 1997. A new genus and species
of lizard (Reptilia: Scincidae) from New Caledonia, southwest
Pacific. Pacific Science 51: 91-96.
Sadlier, R. A., and A. M. Bauer. 1999a. The scincid lizard genus
Sigaloseps (Reptilia: Scincidae) from New Caledonia in the
southwest Pacific: description of a new species and review of the
biology, distribution, and morphology of Sigaloseps deplanchei
(Bavay). Records of the Australian Museum 51(1): 83-91.
http://dx.doi.org/10.3853/j.0067-1975.51.1999.1296
Sadlier, R. A., and A. M. Bauer. 1999b. The scincid lizard Lioscincus
tillieri (Reptilia: Scincidae) from New Caledonia in the southwest
Pacific: new information on the species, biology, distribution, and
morphology. Records of the Australian Museum 51(1): 93-98.
http://dx.doi.org/10.3853/j.0067-1975.51.1999.1297
Sadlier, R. A., and A. M. Bauer. 2002a. The scincid lizard
Graciliscincus shonae (Lacertilia: Scincidae) from New
Caledonia in the southwest Pacific: a review of the species’
morphology, distribution and conservation status. In Zoologia
Neocaledonica 5, ed. J. Najt and P. Grandcolas. Memoires du
Museum national d’Histoire naturelle 187: 257-267.
Sadlier, R. A., A. M. Bauer, and A. H. Whitaker. 2002b. The scincid
lizard genus Nannoscincus Gunther from New Caledonia in the
southwest Pacific: a review of the morphology and distribution of
species in the Nannoscincus mariei species group, including the
description of three new species from Province Nord. In Zoologia
Neocaledonica 5, ed. J. Najt and P. Grandcolas. Memoires du
Museum national d’Histoire naturelle 187: 233-255.
Sadlier, R. A., A. M. Bauer, A. H. Whitaker, and S. A. Smith. 2004a.
Two new species of scincid lizards (Squamata) from the Massif de
Kopeto, New Caledonia. Proceedings ofthe California Academy
of Sciences 55: 208-221.
Sadlier, R. A., A. M. Bauer, and S. A. Smith. 2006a. A new species
of Nannoscincus Gunther (Squamata: Scincidae) from high
elevation forest in southern New Caledonia. Records of the
Australian Museum 58(1): 29-36.
http://dx.doi.org/10.3853/j. 0067-1975.58.2006.1458
Sadlier, R. A., A. M. Bauer, P. L. Wood Jr, S. A. Smith, A. H. Whitaker,
H. Jourdan, and T. Jackman. 2014a. Localized endemism in the
southern ultramafic bio-region of New Caledonia as evidenced
by the lizards in the genus Sigaloseps (Reptilia: Scincidae), with
descriptions of four new species. In Zoologia Neocaledonica
8, ed. E. Guilbert, T. Robillard, H. Jourdan, and P. Grandcolas.
Memoires du Museum national d ’Histoire naturelle 206:79-113.
Sadlier, R. A., A. M. Bauer, S. A. Smith, G. M. Shea, and A. H.
Whitaker, 2014b. High elevation endemism on New Caledonia’s
ultramafic peaks—a new genus and two new species of scincid
lizard. In Zoologia Neocaledonica 8, ed. E. Guilbert, T.
Robillard, H. Jourdan, and P. Grandcolas. Memoires du Museum
national d’Histoire naturelle 206: 115-125.
Sadlier, R. A., G. M. Shea, and A. M. Bauer. 1997. A new genus and
species of lizard (Squamata, Scincidae) from New Caledonia,
Southwest Pacific. In Zoologia Neocaledonica 4, ed. J. Najt, and
L. Matile. Memoires du Museum national d’Histoire naturelle
171: 379-385.
Sadlier, R. A., S. A. Smith, A. M. Bauer, and A. H. Whitaker. 2004b.
Anew genus and species of live-bearing scincid lizard (Reptilia:
Scincidae) from New Caledonia. Journal of Herpetology 38:
320-330.
http://dx.doi.org/10.1670/196-03A
Sadlier, R. A., S. A. Smith, and A. M. Bauer. 2006b. A new genus
for the New Caledonian scincid lizard Lygosoma euryotis
Wemer 1909, and the description of a new species. Records of
the Australian Museum 58(1): 19-28.
http://dx.doi.org/10.3853/j.0067-1975.58.2006.1457
Sadlier, R. A., S. A. Smith, A. M. Bauer, and A. H. Whitaker.
2009. Three new species of skink in the genus Marmorosphax
Sadlier (Squamata: Scincidae) from New Caledonia. In Zoologia
Neocaledonica 7, ed. P. Grandcolas. Memoires du Museum
national d’Histoire naturelle 198: 373-390.
Sadlier, R. A., A. H. Whitaker, and A. M. Bauer. 1998. Lioscincus
marina, a new species of lizard (Reptilia: Scincidae) from New
Caledonia, southwest Pacific. Pacific Science 52: 334-341.
Schmitz, A., I. Ineich, andL. Chirio. 2005. Molecular review of the
genus Panaspis sensu lato (Reptilia: Scincidae) in Cameroon,
with special reference to the status of the proposed subgenera.
Zootaxa 863: 1-28.
Skinner, A., A. F. Hugall, andM. N. Hutchinson. 2011. Lygosomine
phylogeny and the origins of Australian scincid lizards. Journal
of Biogeography 38:1044-1058.
http://dx.doi.org/10.1111 /j. 1365-2699.2010.02471 .x
Skinner, A., M. N. Hutchinson, and M. S. Lee. 2013. Phylogeny
and divergence times of Australian Sphenomorphus group
skinks (Scincidae, Squamata). Molecular Phylogenetics and
Evolution 69: 906-918.
http://dx.doi.org/10.1016/j.ympev.2013.06.014
Smith, M. A. 1937. A review of the genus Lygosoma (Scincidae:
Reptilia) and its allies. Records of the Indian Museum 39:213-234.
Smith, S. A., R. A. Sadlier, A. M. Bauer, C. C. Austin, and T.
Jackman. 2007. Molecular phylogeny of the scincid lizards of
New Caledonia and adjacent areas: evidence for a single origin
of the endemic skinks of Tasmantis. Molecular Phylogenetics
and Evolution 43: 1151-1166.
http://dx.doi.org/10.1016/j.ympev.2007.02.007
Werner, F. 1909. Neue oder seltene Reptilien des Musee Royal
d’Histoire naturelle de Belgique in Brussel. Zoologische
Jahrbiicher 28: 263-288.
Whitaker, A. H., R. A. Sadlier, A. M. Bauer, and V. A. Whitaker.
2005. Biodiversity and Conservation Status of Lizards in Dry
Forest Remnants in Province Sud, New Caledonia. Report by
Whitaker Consultants Limited to Direction des Ressources
Naturelles, Province Sud, Noumea, iv + 59pp.
Whitaker, A. H., and V. A. Whitaker. 2006. Survey of the Lizard
Fauna of the Summit of Massif de Kopeto, Province Nord,
New Caledonia. Unpublished report by Whitaker Consultants
Limited, Motueka to Societe le Nickel, Noumea. 27 pp.
Whiting, A. S., A. M. Bauer, and J. W. Sites. 2003. Phylogenetic
relationships and limb loss in sub-Saharan African scincine
lizards (Squamata: Scincidae). Molecular Phylogenetics and
Evolution 29: 582-598.
http://dx.doi.org/10.1016/SI 055-7903(03)00142-8
Zug, G. R. 2013. Reptiles and Amphibians of the Pacific Islands.
Berkeley: University of California Press. 306 pp.
Manuscript submitted 2 July 2015, revised 6 November 2015, and accepted
9 November 2015.
Records of the Australian Museum
ISSN 0067-1975 (print) ■ ISSN 2201-4349 (online)
http://dx.doi.org/10.3853/issn.2201-4349
Volume 64 (2012) was the last volume with section-sewn saddle-stitching and offset-printing. Improve¬
ments in digital printing allowed us, in 2013, to transition to a high quality, rapid-throughput publication
model with very positive advantages for authors. The new production model also allows publication of
one article per issue with simple saddle-wire binding. Consequently, from Volume 65 (2013), issue size
has decreased and issue number has increased. Binders should note that while we continue to publish one
volume per year we are now releasing a variable number of issues annually.
Records of the Australian Museum continues to be a print and online scientific serial title, with freely
accessible full-text PDF conforming to the most generous Open Access interpretation. The Australian
Museum makes knowledge derived from our collections freely available for all.
The cover of Volume 68 (2016) will be printed with the Australian Museum’s new colours—inspired by
the natural vibrancy of the Australian Red Centre, the desert sand tracks and paths of exploration and
scientific enquiry.
Rec. Aust. Mus. 65(1): 1-37.12 Jun 2013
Rec. Aust. Mus. 65(2): 39-50.24 Jul 2013
Rec. Aust. Mus. 65(3): 51-63.18 Dec 2013
Rec. Aust. Mus. 66(1): 1-62.26 Feb 2014
Rec. Aust. Mus. 66(2): 63-196.2 Apr 2014
Rec. Aust. Mus. 66(3): 197-216.16 Apr 2014
Rec. Aust. Mus. 66(4): 217-240.28 May 2014
Rec. Aust. Mus. 66(5): 241-270.22 Oct 2014
Rec. Aust. Mus. 67(1): 1-24.11 Feb 2015
Rec. Aust. Mus. 67(2): 25-66.13 May 2015
Rec. Aust. Mus. 67(3): 67-108.27 May 2015
Rec. Aust. Mus. 67(4): 109-128.15 Jul 2015
Rec. Aust. Mus. 67(5): 129-161 .7 Oct 2015
Rec. Aust. Mus. 67(6): 163-206.25 Nov 2015
Rec. Aust. Mus. 67(7): 207-224.2 Dec 2015
Records of the Australian Museum — Table of Contents (2013-2015)
CONTENTS
Volume 65 (2013) - Volume 67 (2015)
Ahyong, Shane T. 2014. Deep-sea squat lobsters of the Munidopsis serricornis complex in the Indo-
West Pacific, with descriptions of six new species (Crustacea: Decapoda: Munidopsidae).
Records of the Australian Museum 66(3): 197-216.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1630
Ahyong, Shane T., Kareen E. Schnabel and Keiji Baba. 2015. Southern high latitude squat lobsters:
Galatheoidea and Chirostyloidea from Macquarie Ridge with description of a new species of
Uroptychus. Records of the Australian Museum 67(4): 109-128.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1640
Baba, Keiji.See under Ahyong etal. 2015. 67(4): 109-128
Batley, Michael, and Tony J. Popic. 2013. Five new species of Leioproctus (Protomorpha) Rayment
(Hymenoptera: Colletidae). Records of the Australian Museum 65(2): 39-50.
http://dx.doi.Org/10.3853/j.2201-4349.65.2013.1597
Bauer, Aaron M.See under Sadlier et al. 2015. 67(7): 207-224
Broken-Brow, Julie.See under C. Hughes et al. 2014. 66(4): 225-232
Bruce, Niel L.See under Martin et al. 2014. 66(4): 233-240
Emery, David L.See under Emery et al. 2015. 67(6): 185-199
Emery, Nathan J., David L. Emery and Lindsay W. Popple. 2015. Redescription of Myopsalta atrata
(Goding and Froggatt) and description of two new species of Myopsalta Moulds (Hemiptera:
Cicadidae) from central western New South Wales. Records of the Australian Museum 67(6):
185-199.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1646
Ewart, Anthony, Max S. Moulds and David C. Marshall. 2015. Arenopsaltria nubivena (Cicadidae:
Cicadinae: Cryptotympanini) from the arid regions of Central Australia and southwest Western
Australia. Records of the Australian Museum 67(6): 163-183.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1643
Gerken, Sarah. 2014. Eleven new species and a new genus of Diastylidae (Crustacea: Cumacea) from
Australia and one new species from Canada. Records of the Australian Museum 66(1): 1-62.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1601
Hamilton, Steve.See under C. Hughes et al. 2014. 66(4): 225-232
Harris, Vernon A. 2014. Porcellidiidae of Australia (Harpacticoida, Copepoda). I. A reassessment of
the European species of Porcellidium. Records of the Australian Museum 66(2): 63-110.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1594
Harris, Vernon A. 2014. Porcellidiidae of Australia (Harpacticoida, Copepoda). II. The importance of
the male antennule in taxonomy. Records of the Australian Museum 66(2): 111-166.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1595
Harris, Vernon A. 2014. Porcellidiidae of Australia (Harpacticoida, Copepoda). III. Synopsis of genera
and species. Records of the Australian Museum 66(2): 167-196.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1596
Hill, Kathy B. R.See under Moulds and Hill. 2015. 67(2): 55-66
Hughes, Catherine, Julie Broken-Brow, Harry Parnaby, Steve Hamilton, and Luke K.-P. Leung. 2014.
Rediscovery of the New Guinea Big-eared Bat Pharotis imogene from Central Province, Papua
New Guinea. Records of the Australian Museum 66(4): 225-232.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1632
Hughes, L. E. 2013. Podoceridae of tropical Australia (Peracarida: Amphipoda). Records of the
Australian Museum 65(1): 1-37.
http://dx.doi.Org/10.3853/j.2201-4349.65.2013.1592
Hughes, L. E. 2015. Ampithoidae and Maeridae amphipods from Timor-Leste (Crustacea: Peracarida).
Records of the Australian Museum 67(3): 83-108.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1644
Leung, Luke K.-P..See under C. Hughes et al. 2014. 66(4): 225-232
Marshall, David C.See under Ewart et al. 2015. 67(6): 163-183
Martin, Melissa B., Niel L. Bruce and Barbara F. Nowak. 2014. Smenispa irregularis (Bleeker, 1857)
(Crustacea: Isopoda: Cymothoidae), a buccal-attaching fish parasite from Australia. Records of
the Australian Museum 66(4): 233-240.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1602
Records of the Australian Museum — Table of Contents (2013-2015)
McAlpine, David K. 2014. Review of the Australian genus Pentachaeta (Diptera: Heleomyzidae), with
descriptions of nine new species. Records of the Australian Museum 66(5): 247-264.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1631
McAlpine, David K. 2015. Signal flies of the genus Bama (Diptera: Platystomatidae) in Papua New
Guinea. Records of the Australian Museum 67(2): 25-53.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1603
McEvey, Shane F., and Michele Schiffer. 2015. New species in the Drosophila ananassae subgroup
from northern Australia, New Guinea and the South Pacific (Diptera: Drosophilidae), with
historical overview. Records of the Australian Museum 67(5): 129-161.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1651
McLean, Claire A., Adnan Moussalli, Steve Sass and Devi Stuart-Fox. 2013. Taxonomic assessment of
the Ctenophorus decresii complex (Reptilia: Agamidae) reveals a new species of dragon lizard
from western New South Wales. Records of the Australian Museum 65(3): 51-63.
http://dx.doi.Org/10.3853/j.2201-4349.65.2013.1600
Moulds, Max S.See under Ewart et al. 2015. 67(6): 163-183
Moulds, Max S., and John C. Olive. 2014. A new species of Tamasa Distant from an unusual cave-like
habitat in Australia (Cicadidae: Cicadinae: Tamasini). Records of the Australian Museum 66(5):
265-270.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1637
Moulds, Max S., and Kathy B. R. Hill. 2015. Phylogeny for the tribe Thophini (Cicadoidea:
Cicadidae) with the description of a new subspecies of Thopha sessiliba Distant from Western
Australia. Records of the Australian Museum 67(2): 55-66.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1634
Moussalli, Adnan.See under McLean et al. 2013. 65(3): 51-63
Nowak, Barbara F.See under Martin et al. 2014. 66(4): 233-240
Olive, John C.See under Moulds and Olive. 2014. 66(5): 265-270
Parnaby, Harry.See under C. Hughes et al. 2014. 66(4): 225-232
Popic, Tony J.See under Batley and Popic. 2013. 65(2): 39-50
Popple, Lindsay W..See under Emery et al. 2015. 67(6): 185-199
Sadlier, Ross A., Aaron M. Bauer, Glenn M. Shea, and Sarah A. Smith. 2015. Taxonomic resolution
to the problem of polyphyly in the New Caledonian scincid lizard genus Lioscincus (Squamata:
Scincidae). Records of the Australian Museum 67(7): 207-224.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1649
Sass, Steve.See under McLean et al. 2013. 65(3): 51-63
Schiffer, Michele.See under McEvey and Schiffer. 2015. 67(5): 129-161
Schnabel, Kareen E.See under Ahyong et al. 2015. 67(4): 109-128
Shea, Glenn M.See under Sadlier et al. 2015. 67(7): 207-224
Smith, Graeme B. 2015. New silverfish taxa from Queensland (Zygentoma: Lepismatidae). Records of
the Australian Museum 67(3): 67-81.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1641
Smith, Sarah A.See under Sadlier et al. 2015. 67(7): 207-224
Stuart-Fox, Devi.See under McLean et al. 2013. 65(3): 51-63
Theischinger, Gunther. 2015. Minipteryx robusta —a new genus and species of micropterous
Limnophilinae (Insecta: Diptera: Tipuloidea: Limoniidae) from Australia. Records of the
Australian Museum 67(6): 201-206.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1650
Zabka, Marek. 2014. Maddisonia —a new jumping spider genus from Australia (Arachnida: Araneae:
Salticidae). Records of the Australian Museum 66(4): 217-223.
http://dx.doi.Org/10.3853/j.2201-4349.66.2014.1599
Zhadan, Anna. 2015. Cossuridae (Annelida: Polychaeta: Sedentaria) from Australian and adjacent
waters: the first faunistic survey. Records of the Australian Museum 67(1): 1-24.
http://dx.doi.Org/10.3853/j.2201-4349.67.2015.1639
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