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Phylogeography and taxonomic revision of the New Zealand cryptic skink (Oligosoma inconspicuum; Reptilia: Scincidae) species complex

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The New Zealand skink fauna is highly diverse and contains numerous cryptic, undescribed or hitherto undiscovered species. We completed a taxonomic revision of the cryptic skink (Oligosoma inconspicuum) species complex using molecular (550 bp of the ND2 mitochondrial gene) and morphological analyses. Four new species are described, with each diagnosable by a range of morphological characters and genetic differentiation from several closely related species: O. inconspicuum (sensu stricto), O. notosaurus, O. maccanni, O. stenotis and O. grande. Oligosoma tekakahu sp. nov. is restricted to Chalky Island in Fiordland, and is most closely related to O. inconspicuum and O. notosaurus. The other three new species are restricted to particular mountainous regions in central and western Otago (O. burganae sp. nov., Lammermoor and Rock and Pillar Ranges; O. toka sp. nov., Nevis Valley; O. repens sp. nov., Eyre Mountains) and are most closely related to O. stenotis and O. grande. We also re-described O. inconspicuum. Two proposed new taxa, the 'Big Bay' skink and 'Mahogany' skink, were found to represent Westland/Fiordland populations of O. inconspicuum rather than distinct taxa. We discuss the evolutionary and phylogeographic implications of cryptic and 'anti-cryptic' species within the O. inconspicuum species complex, and suggest that morphologically aberrant populations are the result of local adaptation to novel selective regimes.
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Accepted by S. Carranza: 12 Jan. 2011; published: 3 Mar. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2782: 133 (2011)
www.mapress.com/zootaxa/Article
1
Phylogeography and taxonomic revision of the New Zealand cryptic skink
(Oligosoma inconspicuum; Reptilia: Scincidae) species complex
DAVID G. CHAPPLE1,2,3,7, TRENT P. BELL4, STEPHANIE N.J. CHAPPLE3,5,
KIMBERLY A. MILLER1,2, CHARLES H. DAUGHERTY2 & GEOFF B. PATTERSON6
1School of Biological Sciences, Monash University, Clayton Victoria 3800, Australia
2Allan Wilson Centre for Molecular Ecology and Evolution, School of Biological Sciences, Victoria University of Wellington, P.O. Box
600, Wellington 6140, New Zealand
3Museum Victoria, Division of Sciences, GPO Box 666, Melbourne Victoria 3001, Australia
4EcoGecko Consultants, 212 Pembroke Rd, Wilton, Wellington, New Zealand
5Department of Zoology, University of Melbourne, Melbourne Victoria 3010, Australia
6149 Mairangi Road, Wilton, Wellington, New Zealand
7Corresponding author. E-mail: David.Chapple@monash.edu
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Molecular analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Species descriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Genus Oligosoma Girard, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
Oligosoma inconspicuum (Patterson & Daugherty, 1990) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Oligosoma tekakahu sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
Oligosoma burganae sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Oligosoma toka sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
Oligosoma repens sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25
Oligosoma notosaurus (Patterson & Daugherty, 1990) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
Reference. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Abstract
The New Zealand skink fauna is highly diverse and contains numerous cryptic, undescribed or hitherto undiscovered spe-
cies. We completed a taxonomic revision of the cryptic skink (Oligosoma inconspicuum) species complex using molecular
(550 bp of the ND2 mitochondrial gene) and morphological analyses. Four new species are described, with each diagnos-
able by a range of morphological characters and genetic differentiation from several closely related species: O. inconspic-
uum (sensu stricto), O. notosaurus, O. maccanni, O. stenotis and O. grande. Oligosoma tekakahu sp. nov. is restricted to
Chalky Island in Fiordland, and is most closely related to O. inconspicuum and O. notosaurus. The other three new species
are restricted to particular mountainous regions in central and western Otago (O. burganae sp. nov., Lammermoor and
Rock and Pillar Ranges; O. toka sp. nov., Nevis Valley; O. repens sp. nov., Eyre Mountains) and are most closely related
to O. stenotis and O. grande. We also re-described O. inconspicuum. Two proposed new taxa, the ‘Big Bay’ skink and
‘Mahogany’ skink, were found to represent Westland/Fiordland populations of O. inconspicuum rather than distinct taxa.
We discuss the evolutionary and phylogeographic implications of cryptic and ‘anti-cryptic’ species within the O. incon-
spicuum species complex, and suggest that morphologically aberrant populations are the result of local adaptation to novel
selective regimes.
Key words: cryptic species, Fiordland, Miocene, mitochondrial DNA, molecular clock, morphology, ND2, Otago, Plio-
cene tectonism, Southland, Stewart Island
CHAPPLE ET AL.
2 · Zootaxa 2782 © 2011 Magnolia Press
Introduction
The study of cryptic species is important within the field of evolutionary biology, because they have the potential to
increase our understanding of fundamental concepts such as the nature of species and the mechanisms of specia-
tion. For example, the existence of cryptic species led Ernst Mayr to question the definition of species based on
morphological distinctiveness and to develop the biological species concept, which instead defines species based
on their reproductive isolation (Mayr 1942). Likewise, Mayr’s observations on morphologically aberrant popula-
tions (which could be thought of as the logical opposite of cryptic species) produced fundamental debate about the
roles of drift and selection on speciation and led Mayr to develop the hypothesis of founder-effect speciation (Mayr
1942, 1954; reviewed in Coyne & Orr 2004). The rapid increase in molecular phylogenetic studies since the intro-
duction of the polymerase chain reaction (PCR) has led to renewed interest in cryptic species (Bickford et al.
2007). However, despite the rapidly increasing number of cryptic species being discovered using molecular meth-
ods, our understanding of the processes that underlie cryptic speciation is poor (reviewed in Bickford et al. 2007).
There has been a recent call not only for continued ‘tallying’ of cryptic species but also for scientific inquiry into
the conditions which cause cryptic species to arise (Saez & Lozano 2005).
The New Zealand skink fauna provides a diverse set of potentially informative cases of cryptic species.
Despite the New Zealand archipelago’s temperate latitude and small land area of just 270,000 m2, there are cur-
rently 33 described species in a single genus, Oligosoma (Chapple et al. 2009). For comparison, Australia, an
acknowledged centre of skink diversity, is nearly 30 times as large, but has only 12 times as many described skink
species (Wilson & Swan 2008). It is believed that numerous species of New Zealand skinks remain undescribed
(10–15+ species; Chapple et al. 2009), due largely to the high incidence of cryptic species, as well as a steadily
increasing number of morphologically unusual skinks of uncertain taxonomic affinity being discovered in isolated
and mountainous regions of the archipelago (Jewell & Tocher 2005; Jewell 2007, 2008; but see Chapple & Hitch-
mough 2009).
The cryptic skink, Oligosoma inconspicuum (Patterson & Daugherty), is so named due to its morphological
similarity to several closely-related species and cryptic behaviour (Patterson & Daugherty 1990). Oligosoma
inconspicuum is a small skink with a maximum snout-vent length (SVL) of 70 mm (Patterson & Daugherty 1990).
Its dorsal colour and pattern is highly variable, ranging from a reddish brown with a vertebral stripe to jet black
with no median stripe, while the ventral surface is usually yellow (Patterson & Daugherty 1990). Oligosoma incon-
spicuum was originally discovered using morphological analyses of scale counts and body measurements, and sub-
sequently allozymes, to identify it within a complex of small brown skinks (recognised at the time as Leiolopisma
nigriplantare maccanni, the New Zealand common skink species complex). Taxonomic work on the L. nigriplan-
tare maccanni complex during the 1990’s (Daugherty et al. 1990; Patterson & Daugherty 1990, 1994; Patterson
1997) resulted in the recognition and description of 7 species: O. inconspicuum, O. maccanni (Patterson & Daugh-
erty) (McCann’s skink), O. polychroma (Patterson & Daugherty) (common skink), O. microlepis Patterson &
Daugherty (small-scaled skink), O. notosaurus (Patterson & Daugherty) (southern skink), O. stenotis (Patterson &
Daugherty) (small-eared skink), and O. longipes Patterson (long-toed skink) (In 1995, Patterson & Daugherty res-
urrected Oligosoma to accommodate all New Zealand Leiolopisma species, and Chapple et al. 2009 expanded the
definition of Oligosoma to include all native skink species in New Zealand). Oligosoma inconspicuum could only
be differentiated morphologically from other species within the former common skink species complex by using
multivariate analysis of morphological characters, and it exhibits substantial morphological overlap with O. noto-
saurus, O. polychroma and O. maccanni (Daugherty et al. 1990; Patterson & Daugherty 1990). However, the con-
cept of O. inconspicuum at that time included the species that we describe here as O. burganae sp. nov. and O. toka
sp. nov..
Oligosoma inconspicuum is found in the southern-most regions of New Zealand (Figure 1). Despite the cold
southern climate, at least a dozen other skink species can also be found in these regions (Gill & Whitaker 2001).
Oligosoma inconspicuum occurs in Otago and Southland (Patterson & Daugherty 1990), and on islands in the
northern Foveaux Strait (Whitaker et al. 2002) (Figure 1). It is sympatric with O. maccanni and O. polychroma,
and morphologically difficult to distinguish from either species (Whitaker et al 2002). Oligosoma inconspicuum is
found from sea-level up to 1700 m, and prefers damp, herb or shrubland habitat over tussock and other grassland
(Patterson & Daugherty 1990; Gill & Whitaker 2001; Whitaker et al. 2002). Although sometimes found in grass-
land (Whitaker et al. 2002), it avoids the rocky outcrops favoured by several other species in the region (e.g. the
grand skink, O. grande (Gray), and the Otago skink, O. otagense (Hardy)) (Patterson 1984).
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NEW ZEALAND CRYPTIC SKINK REVISION
FIGURE 1. Map showing the sampling localities for the Oligosoma inconspicuum species complex samples listed in Table 1
(O. inconspicuum, solid black dots; O. notosaurus, white dots; ‘Te Kakahu’ skink [O. tekakahu sp. nov.], white triangles). The
geographical distribution of the new taxa (O. burganae sp. nov., O. repens sp. nov., O. toka sp. nov.) identified in Figure 2 is
shown. Inset: Sampling region, with New Zealand geographic region names used in the text.
The regions of New Zealand in which the species is found are, like the rest of the South Island, geologically
unstable and topographically young. The rugged terrain of the island dates back no further than the Pliocene (Gage
1980; Landis et al. 2008; Wallis & Trewick 2009), and in Otago, some mountain ranges were formed as recently as
the Pleistocene (Jackson et al. 1996). The Southern Alps, which bisect the South Island south-west to north-east
and, rising to over 3000 m, form the South Island’s principal biogeographic barrier, were created by Pliocene uplift
which is still continuing (Gage 1980; Lee et al. 2001; Wallis & Trewick 2009). Prior to this, throughout the Mio-
cene, the island was an eroded peneplain (Cooper & Millener 1993; Cooper & Cooper 1995). The coastlines of the
CHAPPLE ET AL.
4 · Zootaxa 2782 © 2011 Magnolia Press
South Island have also changed radically throughout this time, due to the tectonism as well as rising and falling sea
levels (Newnham et al. 1999). Thus, for example, many modern-day offshore islands within the New Zealand
archipelago have been repeatedly joined to and separated from the mainland. Changing topography and coastlines
are both likely to have implications for the genetic patterns within O. inconspicuum.
Over the past decade there have been suggestions that O. inconspicuum could contain one or more cryptic spe-
cies, since different forms of the species have been observed occurring sympatrically or allopatrically on certain
mountain ranges in Otago (Jewell 2006, 2008). Oligosoma inconspicuum is also thought to harbour ‘anti-cryptic’
species, a term recently coined to describe the situation in which rapid morphological divergence occurs without
substantial genetic change or speciation (Bickford et al. 2007). In this respect, several morphologically unusual
individuals or populations have been discovered in recent years, which are thought to have close affinities with O.
inconspciuum: the ‘Te Kahahu’ skink, the ‘Big Bay’ skink, the ‘Okuru’ skink, and the ‘mahogany’ skink (Jewell
2008). The ‘Te Kakahu’ skink occurs only on Chalky (Te Kakahu) Island off southern Fiordland (Hitchmough et
al. 2007; Jewell 2008; Figure 1). The ‘Big Bay’ skink occurs in large populations in Big Bay, Barn Bay and on the
Cascade Plateau in southern Westland (Miller 1999; Tocher 1999; Jewell 2008; Figure 1). The ‘mahogany’ skink
(Jewell 2008), is known from several specimens and a single tissue sample (Table 1), and was found in the Sinbad
Valley near Milford Sound (see Bell & Patterson 2008 for a detailed site description) (Figure 1). Each taxon has
been proposed to represent a new species (Jewell 2008), but no genetic analysis has been conducted, except for the
‘Big Bay’ skink, where mitochondrial and nuclear DNA sequence data indicates that it is conspecific with O.
inconspicuum (Chapple et al. 2009). Oligosoma inconspicuum is most closely related to O. notosaurus and O. mac-
canni (Chapple et al. 2009), and collectively O. inconspicuum, O. notosaurus and the proposed new taxa are
referred to as the O. inconspicuum species complex.
What processes could underlie the presence of both cryptic and anti-cryptic lineages within the O. inconspic-
uum species complex? Here we use a combination of mitochondrial DNA sequence data (ND2) and morphological
analyses to resolve the numerous taxonomic issues within the O. inconspicuum species complex, and describe new
species where appropriate. We then examine the phylogeographic pattern within the complex, to elucidate the dif-
ferent biogeographic processes which underlie these contrasting (cryptic and anti-cryptic) patterns within O. incon-
spicuum.
Material and methods
Molecular analyses. Samples were obtained from the National Frozen Tissue Collection (NFTC; Victoria Univer-
sity of Wellington, New Zealand) and from ethanol-preserved museum species from Te Papa (National Museum of
New Zealand, Wellington), for sites covering the entire known range of O. inconspicuum and O. notosaurus (Fig-
ure 1, Table 1). Our samples incorporated most of the specimens examined in the descriptions of O. inconspicuum
and O. notosaurus (Patterson & Daugherty 1990), including samples collected at the same locality (Tree Island in
Lake Wakatipu) as the holotype for O. inconspicuum (Table 1). We obtained all of the tissue samples that are avail-
able for the ‘Big Bay’ skink, ‘Te Kakahu’ skink, and the ‘mahogany’ skink (Table 1). However, we were unable to
obtain a tissue sample for the ‘Okuru’ skink. The O. inconspicuum species complex is most closely related to O.
maccanni (Chapple et al. 2009), therefore we included two O. maccanni samples (EF081195, EF447117). We also
included samples from O. stenotis (EU567718, EU567719) and O. grande (EU567720, EU567721), which repre-
sent a well-supported lineage within the New Zealand skink radiation (Chapple et al. 2009), as they are the most
closely-related to the divergent new taxa identified in this study. Two endemic New Caledonian skink species,
Nannoscincus mariei Bavay (earless dwarf skink; EU423132) and Marmorosphax tricolor Bavay (marble-throated
skink; EU423133), were chosen as outgroups based on the broader phylogenetic study of New Zealand skinks
(Chapple et al. 2009).
We extracted total genomic DNA using a modified phenol and chloroform protocol (Sambrook et al. 1989) fol-
lowed by ethanol precipitation. We used PCR with the primers L4437 (5’-AAGCTTTCGGGCCCATACC-3’;
Macey et al. 1997) and ND2r102 (5’- CAGCCTAGGTGGGCGATTG-3’; Sadlier et al. 2004) to amplify a 600-bp
fragment of the ND2 mitochondrial gene. We targeted this region because our previous studies have shown it to be
phylogenetically informative at both the intra- and inter-specific level among New Zealand skinks (Greaves et al.
2007, 2008; Chapple & Patterson 2007; Hare et al. 2008; Liggins et al. 2008a,b; O’Neill et al. 2008; Chapple et al.
2008a,b,c; Miller et al. 2009). PCR and sequencing were conducted as outlined in Greaves et al. (2007).
Zootaxa 2782 © 2011 Magnolia Press · 5
NEW ZEALAND CRYPTIC SKINK REVISION
CHAPPLE ET AL.
6 · Zootaxa 2782 © 2011 Magnolia Press
Sequence data were edited manually using CONTIG EXPRESS in VECTOR NTI ADVANCE 9.1.0 (Invitrogen,
Carlsbad CA, USA) or GENEIOUS Pro 4.8 (Drummond et al. 2010), and trimmed to 550bp. The dataset was then
aligned using CLUSTALX (Thompson et al. 1997) executed in MEGA 4 (Tamura et al. 2007). We translated all
sequences to confirm that none contained premature stop codons. Sequence data were submitted to GenBank under
the accession numbers provided in Table 1. We constructed a TrN neighbour-joining distance phylogram with 1000
bootstraps in MEGA. We considered branches supported by bootstrap values of 70% or greater (Hillis & Bull
1993) to be supported by our data.
To estimate the time since the divergence of lineages within the O. inconspicuum species complex, we cali-
brated the evolutionary rate of ND2 by re-analysing data from Macey et al. (1998) for the agamid genus Laudakia.
Macey et al. (1998) calibrated this rate by geological dating of tectonic events on the Iranian Plateau. Their rate (~
1.2–1.4% per myr) has been demonstrated to be consistent across several vertebrate groups (fish, amphibians, rep-
tiles; reviewed in Weisrock et al. 2001). Specifically, we re-calculated the evolutionary rate for Laudakia using
only the 550bp fragment of ND2 used in the present study (e.g. Smith et al. 2007). We calculated average between-
group nucleotide differences (uncorrected) across each of the calibrated nodes from Macey et al. (1998) (1.5, 2.5
and 3.5 mya), plotted them against time and used the slope of the linear regression to calculate a rate of evolution
for our 550bp fragment of ND2. This resulted in a divergence rate of 1.4% per myr (0.7% evolution per lineage, per
myr) and is slightly faster than the rate of 1.3% per myr found by Macey et al. (1998) for their longer fragment of
ND2.
Morphological analyses. We conducted morphological analyses on all taxa within the O. inconspicuum spe-
cies group (O. inconspicuum, O. notosaurus, ‘Te Kahahu’ skink, ‘Big Bay’ skink, and the ‘mahogany’ skink),
except for the ‘Okuru’ skink. Our analyses built upon the morphological measurements for O. inconspicuum and O.
notosaurus published previously in Patterson & Daugherty (1990). Additional morphological comparisons were
made with O. maccanni (41 specimens) and O. polychroma (72 specimens) (data for both species was obtained
from Patterson & Daugherty 1990). The majority of specimens examined were obtained from Te Papa Tongarewa,
National Museum of New Zealand (RE codes). Several additional specimens (including some holotypes) were col-
lected using the methods outlined in Bell & Patterson (2008), and were lodged at Te Papa.
Descriptions of morphology follow the techniques described in Patterson & Daugherty (1990) and Chapple et
al. (2008a). Midbody scale rows were counted at the midpoint between the fore- and hind legs. Ventral scales were
counted in a line from the mental scale to the vent (including the mental and one preanal scale). The subdigital
lamellae were counted on the fourth hind toe of the right foot. The number of selected head scales were counted
and their arrangement described as outlined in Patterson & Daugherty (1990). Ten measurements (mm) were made
on all specimens: i) axilla to groin (AG), ii) snout to axilla (SF), iii) snout to ear (SE), iv) ear to axilla (EF), head
length (HL) from the posterior part of the interparietal to the tip of the snout, v) head width (HW) between the lat-
eral edges of the left and right parietals, vi) intact tail length (TL), vii) fourth hind toe length from base of toe to tip
excluding nail (FTL), viii) snout-vent length (SVL), ix) hindlimb length (HLL), measured from groin to tip of
fourth toe including nail, and x) forelimb length (FLL), measured from forelimb insertion to tip of fourth finger,
including nail (Patterson & Daugherty 1990, 1994; Chapple et al. 2008a). Numbers are provided where specimens
had missing toes. Colour descriptions are based on the NBS/IBCC colour system (Mundle 1995; see http://
www.anthus.com/Colors/NBS.html).
Results
Molecular analyses
The final dataset contained sequences from 71 individuals in the O. inconspicuum species complex, along with
sequences from three closely-related species (O. maccanni, O. stenotis, O. grande) and two outgroup species. The
final alignment comprised 550 characters from the ND2 mitochondrial gene, of which 228 (41%) were variable and
194 (35%) were parsimony-informative. For the ingroup only, the alignment contained 166 (30%) variable charac-
ters, of which 150 (27%) were parsimony-informative. Base frequencies were unequal (A = 0.316, T = 0.241, C =
0.306, G = 0.137). A 2 test confirmed the homogeneity of base frequencies among sequences (df = 234, P = 1.0).
All sequences were full-length, except that for one sample (OIN38) only 522bp of sequence data was obtained due
to the poor quality of the DNA template.
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NEW ZEALAND CRYPTIC SKINK REVISION
Our molecular analyses indicate that the O. inconspicuum species complex comprises six distinct species (Fig-
ure 1). The new species are diagnosable under both the phylogenetic and morphological species concepts. The first
well-supported taxon (90 bootstrap [BS]) encompasses the type locality for O. inconspicuum (OIN31-36: Tree
Island, Lake Wakatipu) and therefore represents O. inconspicuum sensu stricto (Figure 2). It occurs throughout the
southern South Island with phylogeographic structure evident among Westland/Fiordland, western Otago/South-
land (including Foveaux Strait), and eastern Otago/Southland (Figures 1 and 2). Two proposed new species, tenta-
tively referred to as the ‘Big Bay’ skink and ‘mahogany’ skink, were not found to be distinct species (indeed, the
‘mahogany’ skink was nested within a clade containing the ‘Big Bay’ skink samples). Instead, they represent pop-
ulations of O. inconspicuum (sensu stricto) in Westland and Fiordland, a range extension for the species (Figures 1
and 2).
The second well-supported taxon (91 BS) represents O. notosaurus and is restricted to Stewart Island (Figures
1 and 2). The third taxon is the ‘Te Kakahu’ skink, described here as O. tekakahu sp. nov. (Figures 1 and 2). At
present, this species is only known from Chalky (Te Kakahu) Island in Fiordland (Figure 1). The remaining three
well-supported taxa are genetically divergent (~15%) from O. inconspicuum (sensu stricto) (Table 2) and each have
restricted distributions within particular mountain ranges in central Otago (Figure 1). The fourth taxon (99 BS) is
restricted to subalpine regions of the Lammermoor and Rock and Pillar Ranges of central Otago (Figures 1 and 2),
and is described here as O. burganae sp. nov. (the Burgan skink). The fifth taxon (99 BS) occurs in the Eyre Moun-
tains of western Otago and exhibits substantial genetic divergence (15.7%) from the O. inconspicuum (sensu
stricto) (e.g. OIN1, 25-30, 44) that occur sympatrically in this region (Figures 1 and 2). It is described here as O.
repens sp. nov., the Eyres skink. The final taxon (99 BS) is restricted to the Nevis Valley of central Otago (Figures
1 and 2), and is described here as O. toka sp. nov. (the Nevis skink).
There is substantial variation in the level of genetic divergence among the six species (Table 2). Divergences
among O. inconspicuum (sensu stricto), O. notosaurus and O. tekakahu sp. nov. are relatively low (4.6–8.1%)
compared to that evident between these three species and the three new range-restricted species (O. burganae sp.
nov., O. repens sp. nov., O. toka sp. nov.) in central and western Otago (14.8–16.3%; Table 2). Intermediate levels
of genetic divergence (9.1–10.7%) are present among the three new species in Otago (Table 2). The three new spe-
cies from Otago are most closely related to two species (O. grande: Otago, O. stenotis: Stewart Island endemic)
that also occur in the southernmost regions of New Zealand (Figures 1 and 2; Table 2).
TABLE 2. Mean uncorrected ND2 genetic distances among the species within the Oligosoma inconspicuum species complex.
Comparison is also made to the species (i.e. O. maccanni, O. stenotis, O. grande) that are most closely related to the O. incon-
spicuum species complex.
O.
inconspicuum O.
notosaurus O.
tekakahu O.
burganae O.
repens O.
toka O.
maccanni O.
stenotis O.
grande
O. inconspicuum -
O. notosaurus 0.046 -
O. tekakahu 0.081 0.058 -
O. burganae 0.158 0.154 0.154 -
O. repens 0.153 0.148 0.163 0.107 -
O. toka 0.152 0.155 0.157 0.104 0.091 -
O. maccanni 0.152 0.147 0.148 0.129 0.124 0.116
O. stenotis 0.150 0.147 0.151 0.110 0.106 0.117 0.108 -
O. grande 0.151 0.149 0.167 0.123 0.102 0.103 0.127 0.115 -
CHAPPLE ET AL.
8 · Zootaxa 2782 © 2011 Magnolia Press
FIGURE 2. Neighbour-joining phylogram for the Oligosoma inconspicuum species complex based on 550 bp of the ND2
mitochondrial gene. Bootstrap values (1000 replicates) are shown. Where no support value is shown, the node is unsupported
(i.e. it is less than 70% bootstrap). Six taxa within the O. inconspciuum species complex are identified.
Zootaxa 2782 © 2011 Magnolia Press · 9
NEW ZEALAND CRYPTIC SKINK REVISION
Species descriptions
Genus Oligosoma Girard, 1857
Oligosoma inconspicuum (Patterson & Daugherty, 1990)
Figure 3
Oligosoma inconspicuum Chapple et al. 2009: 485
Oligosoma inconspicuum Jewell 2008: 88
Oligosoma sp. ‘mahogany skink’ Bell & Patterson 2008: 65
Oligosoma sp. ‘mahogany skink’ Bell et al. 2008: 12
Oligosoma sp. 7 (Big Bay skink) Jewell 2008: 90
Oligosoma sp. 8 (mahogany skink) Jewell 2008: 91
Oligosoma inconspicuum Gill & Whitaker 2001: 72
Leiolopisma inconspicuum Patterson & Daugherty 1990: 66
Holotype. Tree Island, Lake Wakatipu (44º 55’S, 168º 25’E), RE002079, adult male (coll. C. Daugherty & R. Mar-
quand, November 1988).
Paratypes (5 specimens). Tree Island, Lake Wakatipu (44º 55’S, 168º 25’E), 4 specimens (RE006135
[CD1905], female; RE006402 [CD1906], male; RE006136 [CD1907], male; RE006137 [CD1908], female) (coll.
I. Southey, March 1986); Tree Island, Lake Wakatipu (44º 55’S, 168º 25’E), RE006169 [FT2063], female (coll.
unknown, December 1988)
Other specimens examined (29 specimens). Gorge Burn, Eyre Mountains (44º 18’S, 168º 15’E), 5 specimens
(RE006127 [CD1101], male; RE006128 [CD1102], female; RE006129 [CD1103], female; RE006013 [CD1104],
male; RE006131 [CD1124], sex unknown) (coll. G. Patterson, March 1985); Eyre Mountains (grid reference
unknown), RE002122, female (coll. I. Southey, April 1986); Macraes Flat (45º 23’S, 170º 26’E), 2 specimens
(RE006141 [CD421], male; RE006142 [CD423], subadult) (coll. C. Daugherty, December 1983); McKenzie
Creek, Big Bay, South Westland (44º 22’S, 168º 02’E), 4 specimens (RE005382 [FT3788], female; RE005383
[FT3789], female; RE005386 [FT3792], female; RE005387 [FT3793], female) (coll. R. van Mierlo & P. van Klink,
January 1998); Barn Bay, Westland (44º 04’S, 168º 18’E), RE005507 [FT3813], male (coll. R. van Mierlo & P. van
Klink, January 1998); Awarua Point, Big Bay, Westland (44º 16’S, 168º 03’E), 4 specimens (RE005434 [FT3031],
female; RE005435 [FT3032], male; RE005436 [FT3033], male; RE005437 [FT3034], male) (coll. I. Southey, Sep-
tember 1992); Awarua Point, Big Bay, Westland (44º 16’S, 168º 03’E), 6 specimens (RE005377 [FT3783], sub-
adult; RE005378 [FT3784], male; RE005371 [FT3785], subadult; RE005380 [FT3786], male; RE005381
[FT3787], female; RE005385 [FT3791], juvenile) (coll. M. Tocher, January 1998); Te Anau township (45º 26’S,
167º 43’E), RE002393, female (coll. G Patterson, February 1985); Tower Peak, Cameron Mountains (46º 01’S,
167º 02’E), RE005497 [FT2924], subadult (coll. G. Gibbs, January 1991); Dipton, Southland (45º 87’S, 168º
23’E), RE001889, female (coll. M. Smith, November 1977); Catlins, Department of Conservation Red Tussock
Reserve (45º 36’S, 167º 14’E), RE006529 [FT3633], male (coll. M. Tocher, January 1997); Sinbad Gully, Llaw-
renny Peaks, Fiordland (44º 38’S, 167º 48’E), RE006880, male (coll. T. Bell February 2008); Mt Nicholas Road,
Eyre Mountains (45º 15’S, 168º 18’E), 4 specimens (RE007284, female; RE007288, female; RE007293, female;
RE007298, female) (coll. J. Reardon, January 2010).
Diagnosis. Oligosoma inconspicuum can be distinguished from other related Oligosoma species through a
combination of characters (Figure 4; Wessa 2011). Compared to O. maccanni, O. inconspicuum has a glossy
appearance, with brown predominating whereas O. maccanni has a greyer ground colour. Oligosoma maccanni has
a pale grey ventral colour rather than the yellow or bronze ventral colour seen in O. inconspicuum. The ear opening
in O. maccanni often has large projecting scales on the interior margin, whereas these are often minimal or lacking
altogether in O. inconspicuum. Longitudinal striping is more pronounced in sympatric populations of O. poly-
chroma, which almost always have a pale stripe on the outside of the forelimbs. The ear opening in O. polychroma
often has prominent projecting scales on the interior margin. There are statistical differences between O. inconspic-
uum and O. burganae sp. nov. (AG/SF, HL/HW, SE/EF, SVL/HL, SVL/FLL), O. notosaurus (SVL/HL, ventral
scales), and O. toka sp. nov (SVL/FLL, SVL/HLL ventral scales) (see Figure 4). Unlike O. repens sp. nov. and O.
toka sp. nov. which have three supraoculars all O. inconspicuum have four supraoculars. Most O. burganae sp.
nov. have only three supraoculars. Oligosoma repens sp. nov. has a more elongate appearance than O. inconspic-
uum (e.g. TL/SVL of 1.28 and 1.16, respectively). The number of subdigital lamellae (17–23) is greater in O.
inconspicuum compared to O. tekakahu sp. nov. (16).
CHAPPLE ET AL.
10 · Zootaxa 2782 © 2011 Magnolia Press
Description of Holotype: Body elongate, oval in cross-section; limbs moderately well-developed, pentadac-
tyl. Lower eyelid with a transparent palpebral disc, bordered on sides and below by small, oblong granules. Nostril
centred just below middle of nasal, pointing up and back, not touching bottom edge of nasal. Supranasals absent.
Rostral broader than deep. Frontonasal broader than long, separated from frontal by scale between prefrontals.
Frontal longer than broad, shorter than frontoparietal and interparietal together, in contact with 2 anteriormost
supraoculars. Supraoculars 4, the second is the largest. Frontoparietals distinct, larger than interparietal. A pair of
parietals meeting behind interparietal and bordered posteriorly by a pair each of nuchals and temporals, also in con-
tact with interparietal, frontoparietal, third/fourth supraocular and 2 postoculars. Loreals 2, anterior one the larger;
anterior loreal in contact with first supralabial, posterior loreal, prefrontal, frontonasal and nasal; posterior loreal in
contact with second supralabial, first subocular, upper and lower preocular, prefrontal and anterior loreal. Suprala-
bials 7, the sixth is the largest. Infralabials 7, several of them equal in size; fifth supralabial below centre of eye.
Mental broader but shallower than rostral. Suboculars 3 and 4 separated by fifth supralabial. Postmental larger than
mental. Chinshields 3 pairs. One primary temporal. Dorsal scales largest, weakly striate. Ventral scales smooth.
Subdigital lamellae smooth. Ear opening round, moderately large, with one projecting granule on anterior margin.
Forelimbs shorter than hindlimbs. Adpressed limbs not meeting in adult. Digits moderately long, cylindrical. Third
front digit shorter than the fourth.
FIGURE 3. Holotype of Oligosoma inconspicuum (RE002079), Tree Island, Lake Wakatipu.
Measurements (in mm; holotype with the variation shown in the paratypes/specimens examined in
parentheses). SVL 61.5 (mean 54.7, range 24.0–74.4), HL 8.5 (mean 7.8, range 5.2–9.7), HW 6.2 (mean 5.4,
range 3.2–6.7), AG 32.1 (mean 28.9, range 10.5–44.9), SF 22.2 (mean 20.1, range 11.0–25.4), SE 10.7 (mean 9.4,
5.7–11.8), EF 11.2 (mean 10.6, range 5.1–14.5), HLL 21.5 (mean 19.1, 11.5–23.7), FLL 15.5 (mean 13.5, 8–16.0)
and TL unknown (not intact) (mean 64.4, range 41.0–82.0; N = 14).
Variation (holotype with the variation shown in the paratypes/specimens examined in parentheses).
Upper ciliaries 7 (mean 7, range 6-9); lower ciliaries 10 (mean 10, range 9–13); nuchals 2 pair (mean 3 pairs, range
1–4 pairs); midbody scale rows 28 (mean 29, range 27–32); ventral scale rows 74 (mean 74, range 67–86); subdig-
ital lamellae 22 (mean 20, range 17–23); supraciliaries 5[right]/6[left] (mean 5, range 5–6); suboculars 6 (mean 7,
range 4–8). Frontonasal not usually separated from frontal by prefrontals meeting in midline. Anterior loreal
always in contact with first and second supralabial, posterior loreal usually in contact with second supralabial only.
Supralabials 7 (usual) or 8, the sixth or seventh the largest. Infralabials 5, 6 (usual), or 7. Third front digit as long as
or shorter than the fourth. Maximum SVL 74.4 mm. Fourteen specimens had intact tails (TL/SVL = 1.16). Ratios
for morphological measurements (± SD): AG/SF: 1.42 ± 0.18; SE/EF: 0.91 ± 0.10; HL/HW: 1.45 ± 0.10.
Zootaxa 2782 © 2011 Magnolia Press · 11
NEW ZEALAND CRYPTIC SKINK REVISION
FIGURE 4 a–b. Notched boxplots comparing the morphological characteristics of the species in the Oligosoma inconspicuum
species group (O. inconspicuum sensu stricto, O. toka sp. nov., O. repens sp. nov., O. notosaurus, O. burganae sp. nov.): a)
axilla-groin(AG)/snout-forelimb (SF), b) snout-vent length (SVL)/head width (HW). Oligosoma tekakahu sp. nov. was not
included in the analysis as only one specimen was available. If there is no overlap between two medians, then the medians are
significantly different at a 95% confidence level.
CHAPPLE ET AL.
12 · Zootaxa 2782 © 2011 Magnolia Press
FIGURE 4 c–d. Notched boxplots comparing the morphological characteristics of the species in the Oligosoma inconspicuum
species group (O. inconspicuum sensu stricto, O. toka sp. nov., O. repens sp. nov., O. notosaurus, O. burganae sp. nov.): c)
snout-vent length (SVL)/forelimb length (FLL), d) ventral scales, e) snout-vent length (SVL)/hind limb length (HLL). Oligo-
soma tekakahu sp. nov. was not included in the analysis as only one specimen was available. If there is no overlap between two
medians, then the medians are significantly different at a 95% confidence level.
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NEW ZEALAND CRYPTIC SKINK REVISION
FIGURE 4 e–f. Notched boxplots comparing the morphological characteristics of the species in the Oligosoma inconspicuum
species group (O. inconspicuum sensu stricto, O. toka sp. nov., O. repens sp. nov., O. notosaurus, O. burganae sp. nov.): e)
snout-vent length (SVL)/hind limb length (HLL), f) snout-vent length (SVL)/head length (HL). Oligosoma tekakahu sp. nov.
was not included in the analysis as only one specimen was available. If there is no overlap between two medians, then the medi-
ans are significantly different at a 95% confidence level.
CHAPPLE ET AL.
14 · Zootaxa 2782 © 2011 Magnolia Press
FIGURE 4 g–h. Notched boxplots comparing the morphological characteristics of the species in the Oligosoma inconspicuum
species group (O. inconspicuum sensu stricto, O. toka sp. nov., O. repens sp. nov., O. notosaurus, O. burganae sp. nov.): g)
head length (HL)/head width (HW), and h) snout-ear (SE)/ear-forelimb (EF). Oligosoma tekakahu sp. nov. was not included in
the analysis as only one specimen was available. If there is no overlap between two medians, then the medians are significantly
different at a 95% confidence level.
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NEW ZEALAND CRYPTIC SKINK REVISION
FIGURE 5. Representative habitats of the members of the Oligosoma inconspicuum species complex. a) O. inconspicuum,
Macraes Flat, Otago. b) O. inconspicuum (mahogany form), Sinbad Valley, Fiordland. c) O. tekakahu sp. nov., Chalky Island,
Fiordland. d) O. burganae sp. nov., Rock and Pillars Range, central Otago. e) O. toka sp. nov. and O. inconspicuum, Nevis Val-
ley, central Otago. f) O. repens sp. nov. and O. inconspicuum, Eyre Mountains, central Otago.
CHAPPLE ET AL.
16 · Zootaxa 2782 © 2011 Magnolia Press
Colouration: Although colouration in the species is extremely variable, brown is the predominant colour. Dor-
sal surface light brown or tan to dark brown, with irregular flecks. Mid-dorsal stripe where present not usually con-
tinuous, and margins not straight. Often a pale dorsolateral stripe extending from near tip of snout to base of tail,
becoming indistinct thereafter. Brown lateral stripe two or more scale rows wide, notched on upper and lower
edges, running from tip of snout through eye towards tip of tail. May contain flecks of dark and light brown. Soles
of feet brown/black. Belly yellow, bronze (in Big Bay animals, Tocher 1999), or deep yellow (in the ‘mahogany’
form), often unmarked. Outer surface of forelimbs brown, speckled with light and dark. Chin and neck may be
pale, usually speckled with black, or uniformly black. There do not appear to be sexually dimorphic colour pat-
terns. Juvenile colouration similar to adult.
Etymology. From inconspicuum (Latin, neuter)—not readily visible, referring both to the difficulty in distin-
guishing this taxon from other similar sympatric species, and the cryptic behaviour of this species in its natural
environment. The common name is the cryptic skink.
Habitat and life history. This species is found throughout the lower South Island, as far north as central Otago
(Figure 5a,b). This species has been recorded throughout the lower South Island and central Otago in the following
Ecological Regions and Districts (McEwen 1987; regions are in capitals, districts in lower case): ASPIRING 51.07
Dart; WAITAKI 64.02 St Mary, 64.04 St Bathans; LAKES 66.03 Richardson, 66.05 Remarkables; CENTRAL
OTAGO 67.01 Lindis, 67.03 Dunstan, 67.04 Maniototo, 67.05 Old Man; LAMMERLAW 68.01 Macraes, CAT-
LINS 70.01 Waipahi; OLIVINE 71.01 Cascade, 71.02 Pyke; FIORD 72.01 Darran; MAVORA 73.02 Eyre, 73.03
Upukerora; WAIKAIA 74.01 Nokomai; GORE 75.01 Gore; SOUTHLAND HILLS 76.01 Takitimu, 76.02 Taring-
atura, 76.03 Hokonui; TE WAE WAE 77.03 Longwood; MAKAREWA 78.01 Southland Plains, 78.02 Waituna.
Similarly, environmental classifications range from (to Level II only): K3; L1, L3, L4; M1; N3, N4, N5, N6; O1;
P5; Q1, Q2, Q3, Q4; R1, R2; and S2 (Leathwick et al. 2003). Consequently this species tolerates an extremely wide
range of environmental conditions from coastal to montane environments in both cold or cool, wet climates and
cool but dry climates with low to moderate radiation (McEwen 1987, Leathwick et al. 2003).
It has a distinct microhabitat preference for herbs and shrubs over tussocks and rocks, and tends to tolerate
quite damp environments such as Sinbad Gully in Fiordland and Big Bay on the West Coast. Adults usually give
birth in February-March, with number of offspring ranging from one to three. Diet consists of fruit and insects (Pat-
terson and Daugherty 1990; Tocher 1999).
Conservation status. Oligosoma inconspicuum is currently considered Not Threatened (Partial Decline) in the
New Zealand Department of Conservation’s national threat classification lists (Hitchmough et al. 2010).
Oligosoma tekakahu sp. nov.
Figure 6
Oligosoma inconspicuum ‘Te Kakahu’ Chapple et al. 2009: 485
Oligosoma sp. 6 (Te Kakahu skink) Jewell 2008: 89
Holotype. Chalky Island (Te Kakahu), Fiordland (46º 03’S, 166º 31’E), RE006879, adult male (coll. T. Bell, Janu-
ary 2008).
Live animals examined. Chalky Island, Fiordland (46º 03’S, 166º 31’E), 16 animals (7 adult males, 6 adult
females, 3 subadults) (data collected by T. Bell, January 2008).
Diagnosis. Oligosoma tekakahu can be distinguished from other related Oligosoma species through a combi-
nation of characters (Figure 4). Compared to O. maccanni, O. tekakahu has a glossy appearance, with brown pre-
dominating whereas O. maccanni has a greyer ground colour. The ear opening in O. maccanni often has large
projecting scales on the interior margin, whereas these are minimal in O. tekakahu. The number of subdigital
lamellae in O. maccanni (19–28) does not overlap with O. tekakahu (16). Longitudinal striping is more pronounced
in adjacent populations of O. polychroma, which almost always have a pale stripe on the outside of the forelimbs
and a prominent mid-dorsal stripe which O. tekakahu lacks. The ear opening in O. polychroma often has prominent
projecting scales on the interior margin. The number of subdigital lamellae in O. tekakahu (16) is less than in any
other member of the O. inconspicuum species complex. The number of ventral scales in O. tekakahu (68) is fewer
than O. toka sp. nov. (70–88). Oligosoma tekakahu has a larger head relative to SVL than any other member of the
O. inconspicuum species complex (its head length, 9.8 mm, was longer than any other specimen measured in the
study).
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NEW ZEALAND CRYPTIC SKINK REVISION
Description of holotype. Body elongate, oval in cross-section; limbs moderately well-developed, pentadactyl.
Lower eyelid with a transparent palpebral disc, bordered on sides and below by small, oblong granules. Nostril cen-
tred just below middle of nasal, pointing up and back, not touching bottom edge of nasal. Supranasals absent. Ros-
tral broader than deep. Frontonasal broader than long, not separated from frontal by prefrontals meeting in midline.
Frontal longer than broad, shorter than frontoparietal and interparietal together, in contact with 2 anteriormost
supraoculars. Supraoculars 3[left]/4[right], the second is the largest. Frontoparietals distinct, larger than interpari-
etal. A pair of parietals meeting behind interparietal and bordered posteriorly by a pair each of nuchals and tempo-
rals, also in contact with interparietal, frontoparietal, third/fourth supraocular and 2 postoculars. Loreals 2, anterior
one the larger; anterior loreal in contact with first supralabial, posterior loreal, prefrontal, frontonasal and nasal;
posterior loreal in contact with second supralabial, first subocular, upper and lower preocular, prefrontal and ante-
rior loreal. Supralabials 7, the sixth is the largest. Infralabials 6, several of them equal in size; fifth supralabial
below centre of eye. Mental broader but shallower than rostral. Suboculars 3 and 4 separated by fifth supralabial.
Postmental larger than mental. Chinshields 3 pairs. One primary temporal. Dorsal scales largest, weakly striate.
Ventral scales smooth. Subdigital lamellae smooth. Ear opening round, small with no projecting granules. Fore-
limbs shorter than hindlimbs. Adpressed limbs not meeting in adult. Digits moderately long, sub-cylindrical. Third
front digit shorter than the fourth.
Measurements (in mm; holotype only). SVL 67.2; HL 9.8; HW 6.8; AG 35.6; SF 24.0; SE 11.3; EF 13.0. TL
unknown (not intact). Ratios for morphological measurements: AG/SF 1.48; SE/EF 0.87; HL/HW 1.44 (in mm;
live animals): The mean SVL size (from 14 live animals) was 64.08 mm (range 50–79 mm), and the mean mass
was 5.6 g (range 1.6–10 g).
Holotype scale counts. Upper ciliaries 7; lower ciliaries 9; nuchals 3 pairs; midbody scale rows 30; ventral
scale rows 73; subdigital lamellae 17; supraciliaries 5; suboculars 6. One primary temporal. Third front digit as
long as the fourth.
Colouration. Dorsal surface light olive to chestnut brown, often with highly irregular random dark flecks. A
broken dorsolateral line formed by a single row of triangular-shaped dark or black flecks runs from the posterior
margin of the eye to the base of the tail. This row is bordered by a pale dorsolateral band approximately a half scale
row wide above, and by a broad dark brown lateral band, below, which commences from the posterior of eye, pass-
ing above limb insertions and concluding near tail base. A pale grey lateroventral band similarly runs from the
lower jaw to the tail base. The edges between the lateral band and lateroventral band is variable in markings in indi-
viduals, from scattered black and white flecks running across the body mid-laterally, to a rudimentary but broken
black above/white below mid-lateral stripes. Ventral surface entirely yellow, or sometimes with pale grey chins.
Ventral surface plain without speckling or other markings. Outer surface of forelimbs strong yellowish brown with
darker and lighter specks. Juvenile colouration is unknown, but likely to be similar to adults.
Etymology. From Te Kakahu, the Maori name for the type locality (= Chalky Island). The common name is the
Te Kakahu skink.
Habitat and life history. Oligosoma tekakahu is currently known only to occupy open coastal herbfield and
prostrate shrubs growing on chalk chip strata at one location on northwestern Chalky Island (Loh 2003; this study;
Figure 5c). Chalky Island is located in the southern part of the FIORD 72.04 Preservation Ecological District
(McEwen 1987), a region which consists of non-glaciated coastal plateaus in the south west (100–500 m asl). Pres-
ervation ED is cool, often cloudy and windy, with high annual rainfall (3200–8000 mm), typical of Environment
O5 (Leathwick et al. 2003). At Chalky Island, these skinks occur in generally high densities at the type location
with over fifty skinks encountered in a small area of 50 x 50 m over two days (this study). The type location is at an
altitude of 134–142 m, at the head of chalk cliffs. The flora around the type location consists of windswept grasses
and complex prostrate shrubs including Olearia avicennifolia, O. oporina, Coprosma spp., Metrosideros umbel-
lata, Phormium cookianum, Coriaria arborea, Helischrysum, Poa colensoi, Haloragis erectus, Griselina littoralis,
Hebe elliptica, Pimelia lyallii, and Chionochloa conspicua (Loh 2003). The skink is currently known to be associ-
ated with Olearia avicennifolia, O. oporina, the grasses Carex spp. and Rytidosperma spp. (this study). Although
the extent of habitat similar to the type location is remarkably small on the entire island, O. tekakahu are also likely
to also occupy the largely inaccessible coastal vegetation and prostate scrub vegetation along the entire coastline of
Chalky Island. Skinks have also adapted to open areas created within the scrub by the cutting of tracks leading to
and away from the type location, an observation also recorded by Loh (2003).
CHAPPLE ET AL.
18 · Zootaxa 2782 © 2011 Magnolia Press
FIGURE 6. a) Holotype of O. tekakahu (RE006879), Chalky Island. b) Lateral view of the head of the O. tekakahu holotype
(RE006879). c) Live specimen of O. tekakahu at the type locality (Chalky Island).
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NEW ZEALAND CRYPTIC SKINK REVISION
Oligosoma tekakahu is a diurnal, terrestrial species, and strongly heliothermic with avid sun-basking behav-
iour even during windy conditions by utilising sheltered micro-sites (Loh 2003, TPB, pers. obs.). The following
observations on Te Kakahu skinks were made by TPB. The skinks will bask readily on chalk flakes in air tempera-
tures up to 23.8 °C, above which they retreat to bask partially shaded within vegetation. Basking extends to as late
as 8 pm in the mid-summer. Oligosoma tekakahu appear to have small home ranges and do not appear to defend
territories, often basking in close proximity to other skinks. Skinks generally forage or bask in close proximity to
cover or vegetation, but will cross quickly between habitat patches over chalk. The sex ratio of captured skinks was
almost equal (8 males:6 females). Almost all adult female skinks were heavily gravid by January, and three neo-
nates were sighted, suggesting the possibility of an unusually early season parturition period in late January. The
smallest reproductively mature individual was a 61 mm SVL gravid female. These skinks flick their tongues rela-
tively frequently as they move. Their tongues were noted to be relatively long and coloured bluish-grey. The inci-
dence of regenerated tails or body scarring evident in the sample (N = 15) was generally high (53%). The New
Zealand falcon (Falco novaezeelandiae) is likely to be a predator and was recorded daily at the type location during
the field work for this study (TPB, pers. obs.). There are no other known sympatric species of lizard on Chalky
Island.
Conservation status. The Te Kakahu skink is currently considered Nationally Critical (criterion A. Naturally
or unnaturally very small natural population; with qualifiers Conservation Dependent, Data Poor, and One Loca-
tion) in the New Zealand Department of Conservation’s national threat classification list (Hitchmough et al. 2010).
These skinks appear to be very locally abundant, and may yet be found in new locations on Chalky Island and
nearby islands such as the Passage and Coal Islands. Protection is provided by the status of a mammalian predator-
free offshore island within a national park, and along with ongoing monitoring for invasive mammal incursions by
the New Zealand Department of Conservation. Potential future threats appear to be the invasive mammal pests (e.g.
rats, Rattus spp., and stoats, Mustela erminea) which are present on the adjacent mainland. These skinks exhibit
highly naive behaviour to potentially novel predators, and would likely suffer severe losses as a result of invasion
of rats or stoats. Stoats were present until 1999 on Te Kakahu Island and the population is likely to be in the process
of recovery from their impacts.
Oligosoma burganae sp. nov.
Figure 7
Leiolopisma inconspicuum Patterson & Daugherty 1990: 66
Leiolopisma nigriplantare maccanni Patterson 1984
Holotype. Burgan Stream, Rock and Pillar Range (45º 35’S, 169º 56’E), RE002390/1, adult female (coll. G. Patter-
son, 1982).
Paratypes (13 specimens). Burgan Stream, Rock and Pillar Range (45º 35’S, 169º 56’E), 12 specimens
(RE006149 [CD765], male; RE006150 [CD766], female; RE006151 [CD767], male; RE006152 [CD768], female;
RE006153 [CD769], female; RE006154 [CD770], male; RE006155 [CD771], male; RE006156 [CD772], female;
RE006157 [CD773], female; RE006159 [CD774], female; RE006160 [CD775], male; RE006161 [CD776],
female) (coll. G. Patterson, April 1984); Burgan Stream, Rock and Pillar Range (45º 35’S, 169º 56’E), RE002391/
1, female (coll. G. Patterson, November 1981).
Diagnosis. Oligosoma burganae can be distinguished from other related Oligosoma species through a combi-
nation of characters (Figure 4). Compared to O. maccanni, O. burganae has a glossy appearance, with brown pre-
dominating whereas O. maccanni has a greyer ground colour. Oligosoma maccanni has a pale grey ventral colour
rather than the yellow/grey ventral colour seen in O. burganae. The ear opening in O. maccanni often has large
projecting scales on the interior margin, whereas these are often minimal or lacking altogether in O. burganae.
Longitudinal striping is more pronounced in sympatric populations of O. polychroma, which almost always have a
pale dorsal stripe on the outside of the forelimbs. The ear opening in O. polychroma often has prominent projecting
scales on the interior margin. There are statistical differences between O. burganae and O. inconspicuum (AG/SF,
HL/HW, SE/EF, SVL/HL, SVL/FLL), O. toka sp. nov. (SVL/HLL, ventral scales), O. notosaurus (ventral scales),
and O. repens sp. nov. (SVL/HL, SVL/FL, AG/SF) (Figure 4). All O. inconspicuum have four supraoculars
whereas most O. burganae have only three supraoculars. Oligosoma repens sp. nov. has a more elongate appear-
CHAPPLE ET AL.
20 · Zootaxa 2782 © 2011 Magnolia Press
ance than O. burganae (TL/SVL of 1.1 and 1.28, respectively). The number of subdigital lamellae (18–23) is
greater than in O. tekakahu (16). The head of O. burganae is noticeably blunter and deeper than O. repens sp. nov.
and O. toka sp. nov. (Figures 4, 7–9).
Description of holotype. Body elongate, oval in cross-section; limbs moderately well-developed, pentadactyl.
Lower eyelid with a transparent palpebral disc, bordered on sides and below by small, oblong granules. Nostril cen-
tred just below middle of nasal, pointing up and back, not touching bottom edge of nasal. Supranasals absent. Ros-
tral broader than deep. Frontonasal broader than long, not separated from frontal by prefrontals meeting in midline.
Frontal longer than broad, shorter than frontoparietal and interparietal together, in contact with 2 anteriormost
supraoculars. Supraoculars 3, the second is the largest. Frontoparietals distinct, larger than interparietal. A pair of
parietals meeting behind interparietal and bordered posteriorly by a pair each of nuchals and temporals, also in con-
tact with interparietal, frontoparietal, third supraocular and 2 postoculars. Loreals 2, anterior one the larger; anterior
loreal in contact with first supralabial, posterior loreal, prefrontal, frontonasal and nasal; posterior loreal in contact
with second supralabial, first subocular, upper and lower preocular, prefrontal and anterior loreal. Supralabials 7,
the sixth is the largest. Infralabials 6, several of them equal in size; fifth supralabial below centre of eye. Mental
broader but shallower than rostral. Suboculars 3 and 4 separated by fifth supralabial. Postmental larger than mental.
Chinshields 3 pairs. One primary temporal. Dorsal scales similar in size to ventral scales, weakly striate. Ventral
scales smooth. Subdigital lamellae smooth. Ear opening round, small with no projecting granules. Forelimbs
shorter than hindlimbs. Adpressed limbs not meeting in adult. Digits moderately long, sub-cylindrical. Third front
digit shorter than the fourth.
Measurements (in mm; holotype with the variation shown in the type series in parentheses). SVL 58.9
(mean 55.0, range 46.8–67.1), HL 8.0 (mean 7.3, range 6.6–9.0), HW 5.4 (mean 5.4, range 4.8–6.3), AG 34.3
(mean 31.8, range 25.2–42.0), SF 20.3 (mean 20.0, range 16.8–23.2), SE 10.0 (mean 8.8, 7.2–10.6), EF 10.3 (mean
10.9, range 8.3–12.9), and TL 56.2.
Variation (holotype with the variation shown in the type series in parentheses). Upper ciliaries 7 (mean 6,
range 5–8); lower ciliaries 10 (mean 9, range 7–10); nuchals 3 pairs (mean 2 pairs, range 0–3 pairs); midbody scale
rows 32 (mean 31, range 30–34); ventral scale rows 76 (mean 75, range 70–82); subdigital lamellae 20 (mean 20,
range 18–23); supraciliaries 5 (mean 6, range 4–6); suboculars 7 (mean 6, range 6–7). Frontonasal never separated
from frontal by prefrontals meeting in midline. Anterior loreal in contact with first or second supralabial. Suprala-
bials 7 (usual) or 8, the sixth or seventh are the largest. Infralabials 6 (usual) or 7. Third front digit as long as
(usual) or shorter than the fourth. Maximum SVL 66.2 mm (shrinkage about 5% based on original records). One
specimen had an intact tail (TL/SVL = 1.00). Ratios for morphological measurements (± SD): AG/SF 1.60 ± 0.17;
SE/EF 0.82 ± 0.09; HL/HW 1.37 ± 0.07. The maximum SVL observed was 65 mm for males and 70 mm for
females. Intact TL/SVL = 1.1 (N=18) (Patterson 1985).
Colouration. Dorsal surface moderate olive to dark olive brown, occasionally black, with irregular flecks. A
median dorsal dark grayish brown longitudinal stripe, 2 half-scale rows wide, well or partially developed, com-
mencing behind the head and passing back to the base of the tail. A light brown dorsal band 2 half-scale rows wide
with light flecks. Another broken dark brown band, 1 half- to 2 half-scale rows wide, shading on to a pale dorsolat-
eral band 1 half- to 2 half-scale rows wide. A pale dorsolateral band, extending from posterior margin of eye to first
one-third of tail. This stripe bordered laterally by a dark brown band usually with notched edges above and below.
A broad dark reddish brown lateral band 1.5 to 2.5 scale rows wide, originating at tip of snout, passing through the
eye and ending near tip of tail, bordered laterally by a very dark brown broken band and with pale scales extending
into it from above and below; sometimes flecked with white. Below this an indistinct pale stripe passes from
beneath the anterior border of the eye through the ear, above the limbs to the tail. This stripe is irregularly defined
below by brown scales which merge gradually with the yellowish grey ventral colouration. Ventral surface usually
speckled with black spots on chin and throat. Outer surface of forelimbs is dark brown with black and white specks.
Juvenile colouration similar to adult, but generally lighter. Ear opening round, small, with no projecting granules
on anterior margin. There do not appear to be sexually dimorphic colour patterns.
Etymology. Refers to the Burgan Stream area, the type locality for the species. The common name is the Bur-
gan skink.
Habitat and life history. Oligosoma burganae appears to be confined to the Rock and Pillar Ranges (Figure
5d) and Lammermoor Ranges (CENTRAL OTAGO: 67.07 Rock and Pillar and LAMMERLAW: 68.02 Waipori;
McEwen 1987) of central Otago, and only occurs above 900 m (i.e. a subalpine species). The Rock and Pillar ED
Zootaxa 2782 © 2011 Magnolia Press · 21
NEW ZEALAND CRYPTIC SKINK REVISION
FIGURE 7. a) Holotype of O. burganae (RE002390/1), Rock and Pillar Range. b) Lateral view of the head of the O. burganae
holotype (RE002390/1). c) Live specimen of O. burganae (photo: J. Reardon).
CHAPPLE ET AL.
22 · Zootaxa 2782 © 2011 Magnolia Press
consists of sub-continental schist block mountains rising steeply from an altitude of 400–1450 m, with annual rain-
fall from 500–1700 mm and snow to 1000 m during winter. This area is classified as Environment Q3 (Leathwick
et al. 2003). The predominant vegetation of the area is montane short and tall Chionochloa tussockland, with some
scrub, particularly Coprosma and Olearia spp. The Waipori ED consists of peaty uplands of the Lammermoor and
Lammerlaw Plataeu (up to 1200m asl), which experience cool, dry or moist subcontintental conditions (annual
rainfall 500–1200 mm), with snow down to 900m during winter. Predominant vegetation is similar to the Rock and
Pillar Ranges, but with pastoral development for sheep and cattle up to 600 m.
The biology, ecology and life-history of O. burganae have been documented previously in Patterson (1985,
1992). The species becomes sexually mature at 49 mm SVL, and has a maximum litter size of six. Parturition
occurs in late January or early February. Some sperm was present in the epididymis from October to March, with a
large increase of sperm in January. The average home range size for O. burganae was 13.4 m2. Specific site defence
was noted on several occasions, usually towards other skink species. The preferred microhabitat appeared to be
herbs and shrubs rather than rocks and grasses. This microhabitat preference appeared to be the main reason why
this species was able to coexist with other similarly-sized skink species (O. polychroma and O. maccanni) through-
out its range. Like most Oligosoma, O. burganae is a diurnal heliotherm. Its diet consists of a range of inverte-
brates, particularly spiders, and berries from several shrub species such as snowberry (Gaultheria depressa). The
climate throughout the species’ range is harsh, where snow may occur at any time of the year. Abdominal fat bod-
ies were noted in many specimens, which increased in size during the summer months, and probably aided the
skinks in hibernation during winter. Tails, likewise, were relatively emaciated after the skinks emerged from hiber-
nation in the spring, and became plumper over summer as the skinks increased their fat reserves. A population cen-
sus from 24 January to 20 February 1983 gave a mean density of one animal per 27 m2 in the type locality. The
survival of O. burganae after a controlled burn-off of an area of tussock grassland was noted by Patterson (1984).
Conservation status. Oligosoma burganae is currently considered At Risk: Declining (B, large population and
low to moderate ongoing or predicted decline; with qualifiers Data Poor, and Range Restricted) in the New Zealand
Department of Conservation’s national threat classification lists (Hitchmough et al. 2010). A recent assessment of
the Burgan skink population (this study) suggests that there has been a serious population decline in the species
since 1985 (Patterson 1985). Thus, research is needed to identify the current population status and trend, establish
the species’ known range, and identify potential threats.
Oligosoma toka sp. nov.
Figure 8
Oligosoma inconspicuum Jewell 2008: 88
Leiolopisma inconspicuum Patterson & Daugherty 1990: 66
Holotype. Schoolhouse Flat, Nevis Valley, (45º 11’S, 168º 59’E), RE007278, adult male (coll. T. Bell, 2009).
Paratypes (12 specimens). Nevis Range F42 5551813.9 2183702.9 (45º 10’S, 168º 52’E), 7 specimens
(RE006165 [CD938], female; RE006166 [CD939], male; RE006163 [CD936], female; RE006164 [CD937],
female; RE006167 [CD940], male; RE006168 [CD941], female; RE006162 [CD935], female) (coll. A.H. Whita-
ker, March 1986); Schoolhouse Flat, Nevis Valley, (45º 11’S, 168º 59’E), 5 specimens (RE007281, female;
RE007286, female; RE007289, female; RE007290, male; RE007297, male) (coll. J. Reardon January 2010).
Live animals examined. Schoolhouse Flat, Nevis Valley (45º 11’S, 168º 59’E), 13 specimens (5 adult males, 5
adult females, 3 juveniles) (data collected by T. Bell, 2009).
Diagnosis. Oligosoma toka can be distinguished from other related Oligosoma species through a combination
of characters (Figure 4). Compared to O. maccanni, O. toka has a glossy appearance, with brown predominating
whereas O. maccanni has a greyer ground colour. Oligosoma maccanni has a pale grey ventral colour rather than
the yellow ventral colour seen in O. toka. The ear opening in O. maccanni often has large projecting scales on the
interior margin, whereas these are often minimal or lacking altogether in O. toka. Oligosoma maccanni has four
supraocular scales compared with three in O. toka, an unusually low number for New Zealand skinks. Sympatric O.
polychroma have very similar colour patterns, but can be distinguished by a pale dorsal stripe on the outside of the
forelimbs, and a greyish-brown ventral colouration. The ear opening in O. polychroma often has prominent project-
Zootaxa 2782 © 2011 Magnolia Press · 23
NEW ZEALAND CRYPTIC SKINK REVISION
ing scales on the interior margin. There are statistical differences between O. toka and O. repens sp. nov. (SVL/HL,
SVL/HLL, ventral scales, SE/EF), O. burganae (SVL/HLL, ventral scales), O. inconspicuum (SVL/FL, SVL/HLL,
ventral scales), and O. notosaurus (ventral scales) (Figure 4). All O. toka have three supraoculars whereas all O.
inconspicuum and O. notosaurus have four. The number of ventral scales in O. tekakahu (68) is fewer than O. toka
(70–88), and the number of subdigital lamellae (16) is fewer than O. toka (17–23). The dorsal surface of the head is
usually more strongly marked than in O. repens sp. nov., and the mid-dorsal and dorsolateral stripes in O. toka are
more prominent than in O. repens sp. nov..
Description of Holotype. Body elongate, oval in cross-section; limbs moderately well-developed, pentadactyl.
Lower eyelid with a transparent palpebral disc, bordered on sides and below by small, oblong granules. Nostril cen-
tred just below middle of nasal, pointing up and back, not touching bottom edge of nasal. Supranasals absent. Ros-
tral broader than deep. Frontonasal broader than long, not separated from frontal by prefrontals meeting in midline.
Frontal longer than broad, shorter than frontoparietal and interparietal together, in contact with 2 anteriormost
supraoculars. Supraoculars 3, the second is the largest. Frontoparietals distinct, larger than interparietal. A pair of
parietals meeting behind interparietal and bordered posteriorly by a pair each of nuchals and temporals, also in con-
tact with interparietal, frontoparietal, third supraocular and 2 postoculars. Loreals 2, similar size; anterior loreal in
contact with first and second supralabial, posterior loreal, prefrontal, frontonasal and nasal; posterior loreal in con-
tact with second supralabial, first subocular, upper and lower preocular, prefrontal and anterior loreal. Supralabials
8[left]/7[right], the sixth and seventh are the equal largest. Infralabials 6, several of them equal in size; sixth[left]/
fifth[right] supralabial below centre of eye. Mental broader but shallower than rostral. Suboculars 3 and 4 separated
by sixth[left]/fifth[right] supralabial. Chinshields 3 pairs. One primary temporal, similar size to lower secondary
temporal. Dorsal scales similar in size to ventral scales, weakly striate. Ventral scales smooth. Subdigital lamellae
smooth. Ear opening round, small with insignificant projecting granules. Forelimbs shorter than hindlimbs.
Adpressed limbs not meeting in adult. Digits moderately long, sub-cylindrical. Third front digit shorter than the
fourth.
Measurements (in mm; holotype with the variation shown in the type series in parentheses). SVL 71.1
(mean 59.8, range 45.1–66.2), HL 8.8 (mean 8.2, range 6.9–9.0), HW 6.4 (mean 5.7, range 4.8–6.5), AG 40.5
(mean 32.0, range 24.1–40.5), SF 24.5 (mean 21.7, range 17.0–24.5), SE 12.2 (mean 10.0, range 8.7–12.2), EF
12.6 (mean 11.7, range 8.3–13.8), and TL 58.5 (mean 58.5, range 53.5–66.0, N=3).
Variation (holotype with the variation shown in the type series in parentheses). Upper ciliaries 7 (mean 6,
range 5–7); lower ciliaries 8 (mean 8, range 7–9); nuchals 4 pairs (mean 3 pairs, range 2–4 pairs); midbody scale
rows 32 (mean 32, range 30–34); ventral scale rows 80 (mean 82, range 70–88); subdigital lamellae 21 (mean 21,
range 17–23); supraciliaries 5 (mean 5, range 5–7); suboculars 7 (mean 6, range 4–7). Frontonasal seldom sepa-
rated from frontal by prefrontals meeting in midline. Anterior loreal in contact with first or second supralabial. Sec-
ondary loreal usually in contact with secondary supralabial only. Supralabials 6, 7 (usual) or 8, the fifth, sixth or
seventh are the largest. Infralabials 5, 6 (usual) or 7. Third front digit as long as (usual) or shorter than the fourth.
Maximum SVL 71.1 mm. Three specimens had an intact tail (TL/SVL = 1.16). Ratios for morphological measure-
ments (± SD): AG/SF 1.47 ± 0.12; SE/EF 0.86 ± 0.11; HL/HW 1.45 ± 0.06.
Colouration. Dorsal surface light to dark yellowish brown, often with irregular flecks. A median dorsal dark
yellowish brown longitudinal stripe, 2 half-scale rows wide, well developed, commencing behind the head and
passing back to the base of the tail, becoming indistinct thereafter. A light to grayish yellowish brown dorsal band
2 half-scale rows wide sometimes with light and dark flecks. This band is often bounded on both sides by a pale
stripe less than one scale wide. Another dark yellowish brown band 1 to 2 half-scale rows wide, shading onto a
prominent pale dorsolateral band 2 half-scale rows wide. This pale band extends from above posterior margin of
eye to base of tail, or further along tail. A broad strong yellowish brown lateral band 1 to 2 scale rows wide, origi-
nating near tip of snout, passing through eye and ending at base or further along tail, bordered laterally by two dark
yellowish brown bands, and often with pale scales extending into it from above and below; sometimes flecked with
white. Below this an indistinct pale stripe passes from beneath the posterior border of the eye above the ear and
limb insertions to the tail. This stripe is irregularly defined below by brown scales which merge gradually with the
yellow ventral colouration. Yellow colouration extends along first third of tail. Ventral surface usually lightly
speckled with black spots on chin and throat, which are white. Outer surface of forelimbs is dark brown with black
and white specks. Juvenile colouration similar to adult, but generally lighter and lacking distinct mid-dorsal stripe.
There do not appear to be sexually dimorphic colour patterns.
CHAPPLE ET AL.
24 · Zootaxa 2782 © 2011 Magnolia Press
FIGURE 8. a) Holotype of O. toka (RE007278), Schoolhouse Flat, Nevis Valley. b) Lateral view of the head of the O. toka
holotype (RE007278). c) Live specimens of O. toka from the type locality.
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NEW ZEALAND CRYPTIC SKINK REVISION
Etymology. From ‘toka’, the Maori word for rock or boulder. Refers to the rocky habitat on which this species
occurs in the Nevis Valley. The common name is the Nevis skink.
Habitat and life history. Oligosoma toka appears to be confined to Nevis Valley (WAIKAIA 74.01 Nokomai
Ecological District; McEwen 1987) of central Otago (Figure 5e). The area where O. toka has been recorded so far
is classified as Environments N3, N4 and Q3 (Leathwick et al. 2003). The Nokomai ED consists of broad plateaus
and hills of a lower altitude (> 600 m to 1500 m asl) than the Eyres ED. The geological composition is complex,
but usually consists of schist and greywacke rock formations. The climate is cool, with annual rainfall typically
around 800–1200 m. It is unclear how far south the Nevis skink population extends to, but it is likely to be the
entire valley and adjacent areas. The predominant ecology consists of exotic pasture for sheep and cattle grazing,
with lowland tussockland in the foothills, and red or subalpine tall tussockland at higher elevations. Oligosoma
toka are extremely abundant around rock piles (old gold tailings) along the eastern side of the Nevis River, but not
as abundant elsewhere where cover is scarce on the open Nevis Valley flats. However, they do occur widely in the
area, on the river flats, around the foothills and up to the Nevis Crossing. The artificial rock piles are likely to be
important refugia for the population in the Nevis Valley, and O. toka is the most abundant species in the valley,
especially adjacent to the Nevis River (T. Jewell, pers. obs.). However, high abundance at certain localities (e.g. 5–
7 skinks per 10 m2 in artificial and natural rock formations; T. Jewell, unpublished data) does not necessarily imply
overall species security, given the apparent restricted range of the species. Important vegetation for O. toka include
tussocks and rank grasses (native, exotic), Coprosma, Discaria Muehlenbeckia, Melicytus and Rubus spp. In the
wider Nevis Valley, O. toka is sympatric with O. inconspicuum, O. polychroma, O. maccanni, Hoplodactylus sp.
‘Cromwell’, and Hoplodactylus sp. ‘Otago large’ (T. Jewell, pers. obs.; this study). It is possible that O. toka might
also be present in the adjacent Hector and Garvie Mountains (T. Jewell, pers. comm.).
Conservation status. Little is known about the range, abundance and population viability of O. toka. It is cur-
rently considered Data Deficient in the New Zealand Department of Conservation’s national threat classification
lists (Hitchmough et al. 2010). Resolution of this species’ conservation status may be urgent (Townsend et al.
2008).
Oligosoma repens sp. nov.
Figure 9
Oligosoma inconspicuum Jewell 2008: 88
Holotype. Mt Nicholas Road, Eyre Mountains, (45º 15’S, 168º 18’E), RE007279, adult male (coll. T. Bell, 2009).
Paratypes (8 specimens). Cascade Creek, Eyre Mountains (45º 13’S, 168º 26’E), 5 specimens (RE007282,
female; RE007285, male; RE007287, male; RE007292, female; RE007294, male) (coll. J. Reardon, January 2010);
Lower Nevis Valley (45º 10’S, 168º 57’E), 3 specimens (RE007291, male; RE007295, male; RE007296, male)
(coll. B. Barratt, December 2009–January 2010).
Diagnosis. Oligosoma repens can be distinguished from other related Oligosoma species through a combina-
tion of characters (Figure 4). Compared to O. maccanni, O. repens has a glossy appearance, with brown predomi-
nating whereas O. maccanni has a greyer ground colour. Oligosoma maccanni has a pale grey ventral colour rather
than the bright yellow ventral colour in O. repens. The ear opening in O. maccanni often has large projecting scales
on the interior margin, whereas these are often minimal or lacking altogether in O. repens. Oligosoma maccanni
has four supraocular scales compared with three in O. repens. Oligosoma polychroma from nearby areas have very
similar colour patterns to O. repens, but can be distinguished by a pale dorsal stripe on the outside of the forelimbs,
and a greyish-brown ventral colouration. The ear opening in O. polychroma often has prominent projecting scales
on the interior margin. There are statistical differences between O. repens and O. toka (SVL/HL, SVL/HLL, ventral
scales), O. burganae (AG/SF, SE/EF, HL/HW, SVL/HL), and O. notosaurus (SVL/HL, ventral scales) (Figure 4).
All O. repens have three supraoculars whereas all O. inconspicuum and O. notosaurus have four. The number of
subdigital lamellae in O. tekakahu (16) is fewer than O. repens (19–23). The dorsal surface of the head is usually
unmarked in O. repens, in contrast with O. toka and O. notosaurus in particular. The species is more gracile than
the other members of the species complex.
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26 · Zootaxa 2782 © 2011 Magnolia Press
FIGURE 9. a) Holotype of O. repens (RE007279), Mt Nicholas Road, Eyre Mountains. b) Lateral view of the head of the O.
repens holotype (RE007279). c) Live specimens of O. repens from the type locality.
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NEW ZEALAND CRYPTIC SKINK REVISION
Description of holotype. Body elongate, oval in cross-section; limbs moderately well-developed, pentadactyl.
Lower eyelid with a transparent palpebral disc, bordered on sides and below by small, oblong granules. Nostril cen-
tred just below middle of nasal, pointing up and back, not touching bottom edge of nasal. Supranasals absent. Ros-
tral broader than deep. Frontonasal broader than long, separated from frontal by prefrontals meeting in midline.
Frontal longer than broad, shorter than frontoparietal and interparietal together, in contact with 2 anteriormost
supraoculars. Supraoculars 3, the second is the largest. Frontoparietals distinct, larger than interparietal. A pair of
parietals meeting behind interparietal and bordered posteriorly by a pair each of nuchals and temporals, also in con-
tact with interparietal, frontoparietal, third supraocular and 2 postoculars. Loreals 2, similar size; anterior loreal in
contact with first and second supralabial, posterior loreal, prefrontal, frontonasal and nasal; posterior loreal in con-
tact with second and third supralabial, first subocular, upper and lower preocular, prefrontal and anterior loreal.
Supralabials 7, the sixth and seventh are the equal largest. Infralabials 6, several of them equal in size; fifth suprala-
bial below centre of eye. Mental broader but shallower than rostral. Suboculars separated by fifth supralabial. Chin-
shields 3 pairs. One primary temporal, approximately half the size of lower secondary temporal. Dorsal scales
similar in size to ventral scales, weakly striate. Ventral scales smooth. Subdigital lamellae smooth. Ear opening
round, small with insignificant projecting granules. Forelimbs shorter than hindlimbs. Adpressed limbs not meeting
in adult. Digits long, sub-cylindrical. Third front digit shorter than the fourth.
Measurements (in mm; holotype with the variation shown in the type series in parentheses). SVL 56.4
(mean 55.1, range 47.6–61.8), HL 8.0 (mean 8.0, range 7.0–9.1), HW 5.7 (mean 5.6, range 4.7–6.0), AG 28.7
(mean 28.8, range 23.6–33.8), SF 20.8 (mean 20.3, range 17.6–22.6), SE 10.5 (mean 10.1, range 9.0–11.3), EF 10.0
(mean 10.4, range 9.1–12.0), and TL unknown (mean 66.5, range 65.0–67.9, N=2).
Variation (holotype with the variation shown in the type series in parentheses). Upper ciliaries 6 (mean 7,
range 5–7); lower ciliaries 10 (mean 9, range 7–10); nuchals 0 pairs (mean 2 pairs, range 0–3 pairs); midbody scale
rows 32 (mean 32, range 30–34); ventral scale rows 77 (mean 76, range 68–81); subdigital lamellae 22 (mean 21,
range 19–23); supraciliaries 6 (mean 6, range 6–7); suboculars 7 (mean 7, range 6–9). Frontonasal seldom sepa-
rated from frontal by prefrontals meeting in midline. Anterior loreal in contact with first or second supralabial. Sec-
ondary loreal usually in contact with secondary and third supralabial. Supralabials 7, the fifth or sixth are the
largest. Infralabials 5 or 6 (usual). Third front digit shorter (usual) or as long as the fourth. Maximum SVL 61.8
mm. Two specimens had intact tails (TL/SVL = 1.28). Ratios for morphological measurements (± SD): AG/SF
1.42 ± 0.07; SE/EF 0.98 ± 0.07; HL/HW 1.43 ± 0.07.
Colouration. Dorsal surface yellowish brown often with a median dorsal very dark brown longitudinal stripe,
2 half-scale rows wide, well or partially developed, commencing behind the head and passing back to the base of
the tail. A yellowish brown dorsal band 2 half-scale to 1.5 scale rows wide sometimes with light flecks. Another
broken dark brown band, 1 half- to 2 half-scale rows wide, shading on to a pale dorsolateral band 1 half- to 2 half-
scale rows wide. This pale dorsolateral band, extending from above and behind posterior margin of eye to base of
tail. This stripe bordered laterally by a strong yellowish brown band 1–2 scale rows wide, originating behind nos-
tril, passing through eye and ending past base of tail, bordered laterally by a dark yellowish brown band. The strong
yellowish band sometimes flecked with white and dark brown. Below this an indistinct pale stripe passes from
beneath the posterior border of the eye through the ear, above the limbs to the base of the tail. This stripe is irregu-
larly defined below by brown scales which merge gradually with the yellow ventral colouration. Ventral surface
may be lightly speckled with black spots on chin and throat, which are white. Outer surface of forelimbs is dark
brown with black and white specks. Juvenile colouration similar to adult, but generally lighter. There do not appear
to be sexually dimorphic colour patterns. Dorsal surface of head normally unmarked.
Etymology. From ‘repens’ (Latin, neuter) = unexpected. Refers to the unexpected discovery of a genetically
divergent new species in the Eyre Mountains that occurs sympatrically with O. inconspicuum (sensu stricto). The
common name is the Eyres skink.
Habitat and life history. The extent of its distribution is unknown, but Oligosoma repens appears to be con-
fined to the Eyre Mountains and also the Hector Mountains (MAVORA 73.02 Eyre; WAIKAIA 74.01 Nokomai;
McEwen 1987) of western Otago (Figure 5f). Environmental classifications for the Eyres are O1 and Q1 and, for
the Hector Mountains, Q1 and Q2 (Leathwick et al. 2003). Oligosoma repens appears to be abundant around rock
piles and screes along the Eyre Mountains foothills (~700 m asl, and likely higher), but less abundant elsewhere
where cover is scarce on the open Eyre valley flats, except where screes occur. The Eyre Ecological District con-
sists of highly dissected, steep and eroding schist or greywacke mountains with narrow valley floors (ranging from
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28 · Zootaxa 2782 © 2011 Magnolia Press
600–2025 m asl) (McEwen 1987). The climate of the Eyres ED is cool and moderately wet (annual rainfall 800
1200 mm); snow may accumulate above 1000 m asl during winter. Much of the Eyres ED was once beech forest,
but have been converted to a mixture of exotic pastoral grasslands and native fescue, red or snow tussockland
(McEwen 1987). The skink is sympatric with O. inconspicuum, O. maccanni, and the gecko Hoplodacylus sp.
‘Otago Large’, another undescribed taxon within the H. maculatus species-complex. Oligosoma chloronoton may
also be a sympatric species (McEwen 1987), as well as O. polychroma.
Conservation status. Little is known about the range, abundance and population viability of O. repens. It is
currently considered Not Threatened (Range Restricted) in the New Zealand Department of Conservation’s
national threat classification lists (Hitchmough et al. 2010). Further research will be required to assess the conser-
vation status of O. repens.
Oligosoma notosaurus (Patterson & Daugherty, 1990)
A full description of O. notosaurus is contained in Patterson & Daugherty (1990). For the current study we re-
examined all of the specimens included in the original description apart from the holotype, which has been lost
from the Te Papa collection.
Discussion
Our taxonomic revision has demonstrated, using molecular and morphological analyses, that the O. inconspicuum
species complex comprises six distinct species and is paraphyletic. While the presence of several undescribed spe-
cies within the O. inconspicuum species complex had been anticipated (e.g. Miller 1999; Tocher 1999; Jewell
2006, 2008; Hitchmough et al. 2007; Chapple et al. 2009), several of our results were surprising. Two of the three
proposed new species (i.e. the ‘Big Bay’ skink and ‘mahogany’ skink) were not found to be distinct species, but
instead represented geographically disjunct populations of the widespread O. inconspicuum sensu stricto. The third
proposed new species (i.e. the ‘Te Kakahu’ skink) was confirmed as a distinct species (O. tekakahu) and was found
to be most closely related (5.8–8.1% genetic divergence) to O. inconspicuum and O. notosaurus. Chapple et al.
(2009) demonstrated that the divergence between O. inconspicuum and O. notosaurus had occurred more recently
compared to most other speciation events within the New Zealand skink radiation. However, the distinct species
status of O. inconspicuum and O. notosaurus is supported by the considerable genetic divergence between these
taxa (4.6%), and the fact that O. notosaurus is not most closely related to the O. inconspicuum populations that
occur on the islands in Foveaux Strait (Figures 1 and 2).
In contrast, the other three new species (O. burganae, O. toka, O. repens) were deeply divergent ‘cryptic’ taxa
(14.8–16.3%) with restricted distributions in the mountainous regions of central Otago. These three species were
found to be more closely related to O. stenotis and O. grande than to the taxon (O. inconspicuum) that they were
previously assigned. This represents a particularly surprising result as O. inconspicuum and its most closely related
species (i.e. members of the former common skink, Leiolopisma nigriplantare maccanni, species complex) have
been the subject of considerable taxonomic study over the past two decades (Daugherty et al. 1990; Patterson &
Daugherty 1990, 1994; Patterson 1997). Neither the allozyme work, nor the discriminate function analyses using
morphological traits, had previously detected these deeply divergent lineages (Daugherty et al. 1990; Patterson &
Daugherty 1990). The cryptic nature of these taxa is highlighted by the fact that specimens from two lineages (i.e.
O. burganae, O. toka) were examined in the original description of O. inconspicuum (Patterson & Daugherty
1990). Indeed, the majority of tissue samples used in this study were obtained from specimens examined in the
original description (Table 1; Patterson & Daugherty 1990).
Our study therefore suggests that there are further cryptic, undescribed or as yet undiscovered members of the
O. inconspicuum species complex present in the southern South Island of New Zealand. The phylogenetic affinities
of the ‘Okuru’ skink are unknown, and since the only known specimen had keeled scales similar to that found in O.
stenotis (Jewell 2008), future work on this taxon may indicate that it also represents a new species.
Intriguingly, several members of the O. inconspicuum species complex occur in sympatry in certain regions of
the South Island. Oligosoma repens occurs sympatrically with O. inconspicuum in at least one locality in the Eyre
Zootaxa 2782 © 2011 Magnolia Press · 29
NEW ZEALAND CRYPTIC SKINK REVISION
Mountains, with substantial genetic divergence (15.7%) between the two species at this location. Similarly, O.
inconspicuum occurs in sympatry with O. toka in the Nevis Valley, the type locality for the latter species (our
observations). In contrast, O. burganae does not occur in sympatry with any other members of the species complex,
but occurs within a few kilometres of an O. inconspicuum population at Macraes Flat (Patterson & Daugherty
1990). However, it does occur together with O. maccanni and O. polychroma in the Rock and Pillars and Lammer-
moor Ranges (Patterson 1984; Patterson & Daugherty 1990). In addition, O. repens also occurs in sympatry with
O. maccanni at several locations (our observations). Oligosoma tekakahu is the only skink species present on
Chalky Island in Fiordland, while O. notosaurus occurs sympatrically with O. stenotis and O. polychroma on Stew-
art Island (Gill & Whitaker 2001; Jewell 2008).
The genetic divergence between the lineage comprising O. inconspicuum, O. notosaurus and O. tekakahu and
the lineage containing O. burganae, O. repens and O. toka was 14.8–16.3%. Given a rate of molecular evolution of
approximately 1.4% per myr, these two lineages are estimated to have diverged during the mid-late Miocene (~10
mya). This corresponds to a period in New Zealand’s history of rapid change, as the climate cooled and dried, and
increases in land area occurred, due to land uplift and falling sea levels (Lee et al. 2001). New habitat formed as
sub-tropical rainforest fragmented and gave way to expanding herb land (Pole et al. 2003), and the newly-activated
Alpine Fault began to form high, rugged hills on what had previously been an eroded peneplain (Cooper & Mil-
lener 1993; Cooper & Cooper 1995; Lee et al. 2001). Novel habitats and range fragmentation during the Miocene
have been implicated in the speciation of other New Zealand endemic taxa. In particular, deeper divergences
among species of giant weta (genus Deinacrida), which includes both alpine and lowland species, have been dated
to the Miocene (Trewick & Morgan-Richards 2005), as has the radiation of alpine cicadas (genus Maoricicada;
Buckley & Simon 2007).
In O. inconspicuum, the genetic distance between the populations in central Otago/Southland and eastern
Otago/Southland is 3.8%, which suggests divergence during the Pliocene (~2.7 mya). Pliocene-age, east-west phy-
logeographic splits across the Otago region have also been observed in the grand skink, O. grande (Berry &
Gleeson 2005), and in McCann’s skink, O. maccanni (O’Neill et al. 2008). This phylogeographic break may be
associated with habitat partitioning either side of the Nevis-Cardrona fault system, which is marked by the Card-
rona and Nevis rivers. The fault marks the boundary between the high relief, mountainous habitat found in the
west, and the undulating grassland habitat found on the lower mountains, hills and wide basins that characterise
central and eastern Otago (Waters et al. 2001).
Oligosoma notosaurus, which is endemic to Stewart Island, is separated from the South Island populations of
O. inconspicuum by Foveaux Strait. Genetic divergence between these two species is 4.6%, which places the split
at 3.3 mya. This is surprising since Foveaux Strait is narrow and relatively shallow. The strait is bridged during gla-
cial periods, most recently ~11,500 years ago when Stewart Island was joined to Southland by a broad coastal plain
(McGlone & Wilson 1996). Evidence for recent (i.e. Pleistocene) geneflow across Foveaux Strait in skinks is
equivocal. In a study of the green skink, O. chloronoton (Hardy), the minimum distance across Foveaux Strait was
similar, at 5.1% (Greaves et al. 2007). However, the common skink, O. polychroma, shows very little structure
across this waterway (Liggins et al. 2008b). This implies that glacial land bridges provided suitable habitat for the
migration or range expansion of some, but not all, species of skink.
In contrast, the genetic distance among the new range-restricted species from central and western Otago (O.
burganae, O. repens and O. toka) is 9.1–10.7%. These three species therefore diverged much earlier than O. incon-
spicuum, O. notosaurus, and O. tekakahu, possibly in the Miocene (6.5–7.6 mya) before the landscape of Otago
became mountainous. Climate change and habitat fragmentation during the Miocene may therefore underlie diver-
gence in these three species (e.g. Lee et al. 2001).
Our study shows that the O. inconspicuum species complex contains both cryptic and ‘anti-cryptic’ taxa (Fig-
ure 10), and raises the question of why both occur within a single complex. The anti-cryptic forms (i.e. ‘Big Bay’
skink, ‘mahogany’ skink) occur in regions of the South Island that are completely isolated, either in valleys on
steep-sided mountains, or in bays surrounded by such mountains. In contrast, O. inconspicuum (sensu stricto)
occurs across a much broader range, and three of the new species (O. burganae, O. repens and O. toka) occur in the
mountainous regions of central and western Otago, sometimes in sympatry with O. inconspicuum. This is an exam-
ple of a geographic pattern noted by Mayr (1942); he observed that peripheral, isolated populations are often mor-
phologically ‘aberrant’, while populations in the interior of a species range show very little morphological
variation. Mayr (1954) reasoned that rapid genetic drift in small, isolated populations, was the main evolutionary
CHAPPLE ET AL.
30 · Zootaxa 2782 © 2011 Magnolia Press
force causing the morphological divergence (and ultimately, speciation) of these peripheral populations. However,
the idea that drift rather than selection drives this geographical pattern has proven controversial (reviewed in Coyne
& Orr 2004). There is little empirical evidence that genetic drift has a major role in morphological evolution
(Coyne et al. 1997) but much demonstrating rapid morphological change in small, isolated populations subjected to
novel selection (reviewed in Turelli et al. 2001). Thus, the unusual anti-cryptic forms in the O. inconspicuum spe-
cies complex are likely to represent local adaptation to the unusual environmental conditions in which these skinks
are found.
FIGURE 10. Comparison of a) the typical form of Oligosoma inconspicuum (top: Macraes Flat, Otago; middle and bottom:
Eyre Mountains, central Otago), and b) the ‘mahogany’ form of O. inconspicuum (Sinbad Valley, Fiordland). The ‘mahogany
form is distinguished by extremely elongated toes, flattened head and deep yellow belly colouration.
Acknowledgements
We thank H. Edmonds, A. Goodman, R. Hitchmough, L. Liggins, P. van Klink, and K. Weston for providing tissue
samples. J. Arrow, R. Cole, R. Coory, H. Edmonds, M. Genet, S. Herbert, R. Hitchmough, T. Jewell, S. Keall, J.
Larivee, E. Loe, C. Miller, D. Morgan, N. Nelson, K. Osborn, J. Reardon, J. Shanks, A. Smart, J. Stahl, P. Thom-
son, M. Tocher, T. Whitaker and C. Wilson provided background information, photographs, logistical support, or
advice during the study. We thank S. Keall, K. Britton and N. Nelson for facilitating access to the frozen tissue col-
lection at VUW, and all of those who collected the samples held in this collection. R. Coory, T. Schultz, S. Whit-
taker and G. Stone provided access to the specimens and tissue samples held in the Museum of New Zealand Te
Papa Tongarewa herpetology collection. We thank Ngai Tahu for supporting this research. The study was funded
by the Allan Wilson Centre for Molecular Ecology and Evolution, and grants from the Department of Conservation
(Conservation Management Units Fund, Investigation No. 4004) to GBP and TPB, Society for Research on
Amphibians and Reptiles in New Zealand (SRARNZ) to DGC and GBP, and Victoria University of Wellington
Research Fund to DGC.
Zootaxa 2782 © 2011 Magnolia Press · 31
NEW ZEALAND CRYPTIC SKINK REVISION
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... Sinbad skinks are medium-sized (to 91 mm SVL) diurnal, cliff-dwelling lizards known from only one site, Sinbad Gully, in the north-western part of Fiordland National Park in the Southland region [15,27], as shown in Figure 2. Mahogany skinks also occur on cliffs in Sinbad Gully (our study site) and are small (to 65 mm SVL) diurnal members of the cryptic skink (O. inconspicuum) species complex [26,27,33]. ...
... Sinbad skinks are medium-sized (to 91 mm SVL) diurnal, cliff-dwelling lizards known from only one site, Sinbad Gully, in the north-western part of Fiordland National Park in the Southland region [15,27], as shown in Figure 2. Mahogany skinks also occur on cliffs in Sinbad Gully (our study site) and are small (to 65 mm SVL) diurnal members of the cryptic skink (O. inconspicuum) species complex [26,27,33]. In evaluating the response of lizards to the drone, we restricted work to fine weather conditions in which lizards were active. ...
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A lack of effective methods for sampling lizards in terrain that is inaccessible to human observers limits our knowledge of their ecology and conservation needs. Drones are increasingly being used in wildlife monitoring, but their potential use for surveying lizards has not been evaluated. We investigated: (1) the detectability of model lizards using a drone relative to a human observer, and (2) the response of four lizard species to an approaching drone in three habitat types. Model lizards placed in potential basking positions within a defined search area were detected by both the drone operator and human observer, but the probability of detection was lower with the drone. Jewelled geckos (Naultinus gemmeus) in shrubland and grand skinks (Oligosoma grande) in rocky habitats showed surprisingly little reaction to the approaching drone, enabling close approaches (means of 59 cm and 107 cm, respectively) and accurate species identification with photos taken by the drone camera. For highly patterned jewelled geckos, identification was also possible on an individual level. However, the drone was unsuccessful at detecting two alpine skink species in a near-vertical cliff habitat. Collectively, our results suggest that drones have potential as a tool for detecting small-bodied lizards in habitats inaccessible to human observers.
... The number of skink species discovered in New Zealand has been steadily growing, and 15 new species have been described in the last 10 years alone (Chapple et al. 2011;Jewell 2017Jewell , 2019Melzer et al. 2017Melzer et al. , 2019Patterson et al. 2013;Patterson & Hitchmough 2021;Whitaker et al. 2018) bringing the total number of recognised species to 50. An additional 15 proposed species have been informally identified, bringing the diversity recognised by van Winkel et al. (2018) to 65 species. ...
... To confirm the distinctiveness of the Bream Head taxon, and determine its phylogenetic position, we sequenced the mitochondrial DNA gene (mtDNA), ND2. Previous studies have found this mtDNA region to be phylogenetically informative for taxonomic and phylogeographic studies of New Zealand skinks (Chapple & Patterson 2007;Chapple et al. 2008aChapple et al. ,b,c, 2009Chapple et al. , 2011Chapple et al. , 2012Greaves et al. 2007Greaves et al. , 2008Hare et al. 2008;Liggins et al. 2008a,b;Melzer et al. 2019;Miller et al. 2009;O'Neill et al. 2008;Patterson et al. 2013;Whitaker et al. 2018). ND2 sequences from the three Bream Head samples were produced by EcoGene Ltd, following the protocols of Greaves et al. (2008), and were compared to previously published ND2 sequences across the range of O. zelandicum (Figure 1; O'Neill et al. 2008). ...
Article
New Zealand is home to a diverse cool temperate assemblage of skinks, with 60+ identified taxa (genus Oligosoma Girard), of which only 50 have been formally described. Here we describe a new species (Oligosoma kakerakau sp. nov.) from Bream Head Scenic Reserve, near Whangārei Heads, Northland. This species is considered to be conspecific with a single specimen (Oligosoma “Whirinaki”) previously reported (in 2003) from Whirinaki Te Pua-a-Tāne Conservation Park ~370 km further south. Oligosoma kakerakau sp. nov. can be distinguished from all other members of the genus by a combination of a distinctive “teardrop” marking below the eye, a distinctive mid-lateral stripe, and the colouration and pattern on its ventral surface. Our phylogenetic analyses indicate that Oligosoma kakerakau sp. nov. is most closely related to O. zelandicum (Gray), and more distantly to O. striatum (Buller) and O. homalonotum (Boulenger). Sea level changes during the Pliocene, such as the formation of the Manawatū Strait, may have contributed to the divergence between Oligosoma kakerakau sp. nov. and O. zelandicum. We discuss the distribution, ecology and conservation of Oligosoma kakerakau sp. nov., and outline future research and conservation priorities for the species.
... New taxa are being identified almost yearly as previously recognised species are redefined as species complexes (e.g. Chapple et al. (2011) or new species are discovered (e.g. Bell & Patterson (2008), Jewell (2017)), including the white-bellied skink Oligosoma hoparatea Whitaker et al. 2018. ...
... We focused on the mitochondrial DNA gene, NADH dehydrogenase 2 (ND2), as previous studies have shown this gene to be phylogenetically informative at both the intra-and inter-specific level among New Zealand skinks, and to produce tree topologies identical to those from larger genetic datasets (Greaves et al. 2007(Greaves et al. , 2008Chapple & Patterson 2007;Hare et al. 2008;Liggins et al. 2008a,b;O'Neill et al. 2008;Chapple et al. 2008aChapple et al. ,b,c, 2009Chapple et al. , 2011Chapple et al. , 2012Miller et al. 2009;Patterson et al. 2013;Whitaker et al. 2018). ND2 sequences from the four Lonely Lake specimens were produced by EcoGene Ltd, following the protocols of Greaves et al. (2008). ...
Article
A new species of Oligosoma is described from a slate scree in montane tussock grassland in Kahurangi National Park, New Zealand, where it is currently known from a single small site. The new species (Oligosoma kahurangi sp. nov.) can be distinguished from all congeners by its extremely long tail, 36–38 mid-body scale rows, head length/head width ratio of 1.66, and colour pattern. It is part of the O. longipes Patterson species complex. The species is currently very poorly known but likely to be highly threatened, and we suggest listing as Nationally Critical (Data Poor, One Location) in New Zealand, and Data Deficient in the IUCN red-list. Predation by introduced mammals, particularly mice, is assumed to be a threat to its survival.
... Likewise, geckos and skinks, which comprise the vast majority of New Zealand's herpetofauna, both experienced their main period of diversification within the Miocene, subsequently radiating into nearly every available habitat type, including the alpine zone (Chapple et al. 2009;Nielsen et al. 2011), and even intraspecific variation among South Island populations often predates the Pleistocene (e.g. Greaves et al. 2007;Chapple et al. 2011;Chapple et al. 2012). Some alpine species may have experienced expanded ranges during the Pleistocene, as suggested by ENMs and high, localised haplotype diversity for the grasshopper Sigaus australis (Carmelet-Rescan et al. 2021), high population structure in the cicada Maoricicada campbelli ), and the persistence of numerous lineages across diverse taxa that are today characterised by relatively small geographic ranges. ...
... This intriguing result drastically conflicts the morphological evidence, which clearly separates the two new species from their respective sympatric congeners. This phenomenon is most likely caused by "anti-cryptic" speciation, where strong divergent selection forces caused rapid morphological divergence (Bickford et al. 2007;Chapple et al. 2011). Other cases that resemble "anti-cryptic" speciation can be found in other freshwater crabs, e.g., Geothelphusa Stimpson, 1858 from Taiwan (the G. olea Shy, Ng & Yu, 1994 complex;Shih et al. 2007a), Somanniathelphusa Bott, 1968(including S. amoyensis Naiyanetr & Dai, 1997, S. taiwanensis Bott, 1968 ...
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Two morphologically distinct new species of Nanhaipotamon Bott, 1968, are described from Huizhou, Guangdong Province, southern China. The smooth carapace and lack of flagellum on the third maxilliped exopod immediately separate N. incendium n. sp. from all other known Nanhaipotamon. It is also the first species of this genus to be recorded from a relatively high altitude of 700 m a.s.l. Nanhaipotamon aureomarginatum n. sp. is externally much more typical of species of its genus but can be distinguished from congeners by its unique male first gonopod. The two new species are more or less sympatric with a morphological disparate congener, which we tentatively assign as N. aff. aculatum Dai, 1997 and N. aff. hongkongense (Shen, 1940), respectively, but with different habitats. A genetic analysis using the mitochondrial cytochrome oxidase subunit I shows that while the two new species are sufficiently distinct from other known species of Nanhaipotamon, they have extraordinarily close relationships with their respective sympatric congeners, which calls for further investigation. The ecology of these two new species are also noted.
... Further recent examples include snipe (Baker et al. 2010), NZ storm petrel , skinks (Chapple et al. 2011;Patterson et al. 2013;Melzer et al. 2017), forget-me-not (Meudt et al. 2013), orchids (Lehnebach et al. 2016) and grasses (de Lange et al. 2016). ...
Article
Few areas of conservation biology have grown at quite the same pace as conservation genetics. New Zealand exemplifies this growth with a 50–100-fold increase in publications since a review in 1994. A wide array of techniques in the fields of population genetics, molecular systematics and molecular biology has now become available to conservation biologists to apply to management. Here I review developments categorised broadly into six headings: measuring diversity among individuals; inbreeding; selection and drift; identification of individuals; measuring differentiation (demes, populations, MUs, ESUs, species, hybrids); and other molecular biological approaches. The vast range of available techniques and analyses makes it more important than ever that appropriate tools are chosen for the questions posed, and applied to where need is greatest, if we are to manage our biological diversity successfully in the twenty-first century.
... For both New Zealand skinks and geckos, the current taxonomy was derived from molecular and morphological studies (reviewed in Chapple & Patterson, 2007;Chapple et al., 2008aChapple et al., ,b, 2009Chapple et al., , 2011Patterson et al., 2013;Chapple & Ritchie, 2013;Hitchmough et al., 2013). For the geckos, many long-recognized and widely accepted species-level forms exist but currently await formal description (Nielsen et al., 2011;Hitchmough et al., unpub. ...
Article
AimConservation is often prioritized by identifying regional clusters of threatened or endemic species. Another approach is to assess the evolutionary distinctiveness of groups of taxa using phylodiversity measures. However, quantification of evolutionary history has traditionally not accounted for its uneven geographical distribution due to the variation in species ranges. We assess the efficacy of phylogenetic endemism (PE) to predict high extinction risk in comparison to estimates of species range restriction (weighted endemism, WE) and phylogenetic diversity (PD). PE measures the relative range restriction of evolutionary history (lineages), while WE concentrates on the tips of the tree of life, treating all such branches as being of equal length. Location/Methods Using New Zealand's endemic skinks and geckos, we mapped the geographical variation in their extinction risk, PE, WE and PD and measured the extent to which extinction risk exhibited phylogenetic clustering for each group. Correlations between geographical concentrations of high skink and gecko extinction risk with PE, WE and PD were calculated. ResultsPE was predictive of spatial clusters of high extinction risk for geckos (r(2)=0.34, P<0.001) while WE was markedly less so (r(2)=0.19, P<0.001). The reverse applied to skinks, with WE most predictive of high risk (r(2)=0.26, P<0.001). The phylogenetic signal of extinction risk was significantly conserved for geckos, but was weaker and non-significant for skinks. PE and WE were not predictive of low risk. PD was not predictive of risk. Main conclusionsPE and related measures may be predictive of extinction risk when risk is phylogenetically conserved. Mapping the geographical variation in PE could be a useful first assessment of extinction risk for many groups because phylogenies are increasingly available, while full risk status categories are not. These findings might apply to other groups and locations and warrant further investigation.
Article
The genera Lepidothyris, Lygosoma and Mochlus comprise the writhing or supple skinks, a group of semi-fossorial, elongate-bodied skinks distributed across the Old World Tropics. Due to their generalized morphology and lack of diagnostic characters, species- and clade-level relationships have long been debated. Recent molecular phylogenetic studies of the group have provided some clarification of species-level relationships, but a number of issues regarding higher level relationships among genera still remain. Here we present a phylogenetic estimate of relationships among species in Lygosoma, Mochlus and Lepidothyris generated by concatenated and species tree analyses of multilocus data using the most extensive taxonomic sampling of the group to date. We also use multivariate statistics to examine species and clade distributions in morpho space. Our results reject the monophyly of Lygosoma s.l., Lygosoma s.s. and Mochlus, which highlights the instability of the current taxonomic classification of the group. We, therefore, revise the taxonomy of the writhing skinks to better reflect the evolutionary history of Lygosoma s.l. by restricting Lygosoma for Southeast Asia, resurrecting the genus Riopa for a clade of Indian and Southeast Asian species, expanding the genus Mochlus to include all African species of writhing skinks and describing a new genus in Southeast Asia.
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New Zealand has a diverse skink fauna, comprising 45 described native species, and at least 15 undescribed taxa, within the single genus Oligosoma Girard, 1857. One of the earliest described, and best known, species is the speckled skink, Oligosoma infrapunctatum (Boulenger 1887). Despite a relatively stable taxonomic history for nearly 114 years, recent molecular work has indicated that O. infrapunctatum represents a species complex, comprising numerous genetically divergent, range restricted taxa. We completed the first stage of a taxonomic revision of O. infrapunctatum, conducting a morphological re-evaluation of existing voucher material, and newly collected specimens, and generated a molecular phylogeny for the species complex. This allowed us to distinguish six species: O. infrapunctatum, two species resurrected from synonymy (O. newmani, O. robinsoni), and three new species (O. salmo sp. nov., O. albornense sp. nov. O. auroraensis sp. nov.). The name bearing type population of O. infrapunctatum has not been located again for at least 130 years: it remains to be rediscovered and may already be extinct. Two of the six species here are considered ‘Nationally Critical’ (O. albornense sp. nov., O. salmo sp. nov.) under the New Zealand Threat Classification System, the others are Nationally Vulnerable (O. auroraensis sp. nov.) and At Risk—Relict (O. newmani, O. robinsoni). Further taxonomic work will be required to determine the taxonomy of other speckled skink genetic lineages in the South Island, particularly O. aff. infrapunctatum “cobble”, O. “Hokitika”, O. “Southern North Island” and O. “Westport”.
Article
New Zealand has a diverse, endemic skink fauna, which is recognised as the most species rich skink assemblage of any cool temperate region on earth. All native New Zealand skink species are assigned to a single genus, Oligosoma Girard. A new species of Oligosoma is described from screes in montane tussock grassland in the mid-Canterbury high country, New Zealand, where it is currently known from four sites on two mountain ranges. The new species (Oligosoma hoparatea sp. nov.) can be distinguished from all congeners by a combination of mid-body scale row and lamellae counts, scale morphologies, and a bold striped pattern with smooth-edged, dark lateral bands. It is part of the O. longipes Patterson species complex, and occurs in sympatry with its closest relative, O. aff. longipes ‘southern’. The species is currently highly threatened, and is listed as Nationally Critical in New Zealand. Predation by a suite of introduced mammals is assumed to be a major threat to its survival.
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A new species of alpine skink, Oligosoma pikitanga, is described from the Llawrenny Peaks, Fiordland, New Zealand. This species is diagnosed on the basis of strong morphological, ecological, and genetic differentiation from the following relatives O. acrinasum, O. infrapunctatum, O. otagense, O. taumakae and O. waimatense. The species is characterized by a shiny black base colour with bright green dorsal blotches, lateral pinkish spots and a vivid orange belly. It appears this new taxon is extremely rare, and at enhanced risk from introduced mammalian predators.
Article
A new species of alpine skink, Oligosoma pikitanga, is described from the Llawrenny Peaks, Fiordland, New Zealand. This species is diagnosed on the basis of strong morphological, ecological, and genetic differentiation from the following relatives O. acrinasum, O. infrapunctatum, O. otagense, O. taumakae and O. waimatense. The species is characterized by a shiny black base colour with bright green dorsal blotches, lateral pinkish spots and a vivid orange belly. It appears this new taxon is extremely rare, and at enhanced risk from introduced mammalian predators.
Article
We evaluate Sewall Wright's three-phase "shifting balance" theory of evolution, examining both the theoretical issues and the relevant data from nature and the laboratory. We conclude that while phases I and II of Wright's theory (the movement of populations from one "adaptive peak" to another via drift and selection) can occur under some conditions, genetic drift is often unnecessary for movement between peaks. Phase III of the shifting balance, in which adaptations spread from particular populations to the entire species, faces two major theoretical obstacles: (1) unlike adaptations favored by simple directional selection, adaptations whose fixation requires some genetic drift are often prevented from spreading by barriers to gene flow; and (2) it is difficult to assemble complex adaptations whose constituent parts arise via peak shifts in different demes. Our review of the data from nature shows that although there is some evidence for individual phases of the shifting balance process, there are few empirical observations explained better by Wright's three-phase mechanism than by simple mass selection. Similarly, artificial selection experiments fail to show that selection in subdivided populations produces greater response than does mass selection in large populations. The complexity of the shifting balance process and the difficulty of establishing that adaptive valleys have been crossed by genetic drift make it impossible to test Wright's claim that adaptations commonly originate by this process. In view of these problems, it seems unreasonable to consider the shifting balance process as an important explanation for the evolution of adaptations.
Article
Analysis of allozyme variation at 17 loci reveals that the endemic New Zealand skink Leiolopisma nigriplantare (sensu Hardy, 1977) consists of a group of at least four cryptic species. One species, redefined as L. nigriplantare, occurs widely in the southern half of the North Island, most of the South Island, and small islands within the Chatham Island group. Leiolopisma maccanni and L. inconspicuum occur in the southern half of the South Island, and L. notosaurus on Stewart Island. A fifth species, L. microlepis, superficially resembles some members of this group and is known from only two small populations in the central North Island. Leiolopisma maccanni and L. microlepis can be identified morphologically, but the other species overlap considerably in morphology and color, explaining the historical problems of resolving this complex of species. The extensive genetic divergence among the species in this complex and also L. zelandicum, whose taxonomic history has been confused with the L. nigriplantare complex, and the high species diversity and endemism of the New Zealand Leiolopisma suggest that Leiolopisma is an ancient New Zealand lineage and has undergone extensive evolution in situ.
Article
We evaluate Sewall Wright's three-phase 'shifting balance' theory of evolution, examining both the theoretical issues and the relevant data from nature and the laboratory. We conclude that while phases I and II of Wright's theory (the movement of populations from one 'adaptive peak' to another via drift and selection) can occur under some conditions, genetic drift is often unnecessary for movement between peaks. Phase III of the shifting balance, in which adaptations spread from particular populations to the entire species, faces two major theoretical obstacles: (1) unlike adaptations favored by simple directional selection, adaptations whose fixation requires some genetic drift are often prevented from spreading by barriers to gene flow; and (2) it is difficult to assemble complex adaptations whose constituent parts arise via peak shifts in different demes. Our review of the data from nature shows that although there is some evidence for individual phases of the shifting balance process, there are few empirical observations explained better by Wright's three-phase mechanism than by simple mass selection. Similarly, artificial selection experiments fail to show that selection in subdivided populations produces greater response than does mass selection in large populations. The complexity of the shifting balance process and the difficulty of establishing that adaptive valleys have been crossed by genetic drift make it impossible to test Wright's claim that adaptations commonly originate by this process. In view of these problems, it seems unreasonable to consider the shifting balance process as an important explanation for the evolution of adaptations.
Article
The modern New Zealand angiosperm flora has many notable characteristics, such as a predominance of evergreen, perennial life forms, few nitrogen-fixing species, despecialised floral features and asymmetric genus—species relations. The origin of these features has been attributed to antiquity of the flora, isolation and/or environmental history. Using evidence from palynology and macrofossils, we investigate the characteristics of the mid–late Cenozoic angiosperm flora and the impact of environmental changes in land area and configuration, physiography and climate on the depletion and composition of the New Zealand flora. Climatic cooling, increasing isolation and tectonism have each acted as important environmental filters, contributing to regional extinctions and decreasing floral diversity, and inducing major turnover in the floristic composition of New Zealand. During the Miocene and Pliocene at least 15 families and a minimum of 36 genera were lost from the New Zealand flora. These included a range of life forms and physiognomically important taxa such as Acacia, Bombax, Casuarina, Eucalyptus, Ilex, many Proteaceae and several palms. The extinction and decline in richness of subtropical families was caused by the onset of cooling conditions in the Late Miocene—Pliocene, and exacerbated by the absence of significant land areas to act as refugia at lower latitudes. Many of these genera/families persist today on islands to the north (e.g. New Caledonia), reflecting mid-Cenozoic land conduits, and in Australia. The close floristic links with New Caledonia were probably maintained by intermittent island stepping-stones which facilitated interchange of subtropical taxa until the Late Miocene. The Pleistocene extinction of some genera, tolerant of warm-temperate environments (e.g. Acacia, Eucalyptus) may be a reflection of the fact that persistent mesic conditions favoured widespread dominance of dense rainforest during interglacials. The loss of these groups, containing diverse life forms and floral structures, suggests that many of the present characteristics of the New Zealand flora reflect strong selective pressures, mainly driven by climate change, in the Late Miocene, Pliocene and Pleistocene, rather than events of greater geological antiquity.