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A new species of Arenophryne (Anura: Myobatrachidae) from the central coast of
Western Australia

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https://escholarship.org/uc/item/7m92h07r

Journal
Records of the Western Australian Museum, 24

ISSN
0312-3162

Authors
Doughty, P
Edwards, DL

Publication Date
2008

Peer reviewed

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Records of the Western Australian Museum 24: 121–131 (2008).

A new species of Arenophryne (Anura: Myobatrachidae)

from the central coast of Western Australia

Paul Doughty1 and Danielle Edwards2

1
Department of Terrestrial Zoology, Western Australian Museum, 49 Kew Street, Welshpool,
Western Australia 6106, Australia. Email: Paul.Doughty@museum.wa.gov.au
2
School of Animal Biology, University of Western Australia,
Crawley, Western Australia 6009, Australia

Abstract – The sandhill frog, Arenophyrne rotunda, belongs to a monotypic

genus that occurs on the central coast of Western Australia. It has a highly
modified body shape with a small head and large front limbs. Members of
this species burrows forwards through sand substrates. Here we describe a
new species of Arenophyrne from the Geraldton sandplain that occurs to the
south of the populations of the type species A. rotunda at Shark Bay. Relative
to A. rotunda, the new taxon has a more pointed snout, smaller face and eyes,
larger hands, rougher dorsal surface and darker colouration that matches the
background colour of the sands on which it occurs. Molecular evidence
indicates divergence of the two taxa in the late Miocene to early Pliocene,
approximately 5–6 mya. The western coast of Australia has a complex
biogeographic history owing to geological activity and changes in sea level
interacting with extensive sandy areas. Speciation within Arenophyrne on the
coastal dunes of Western Australia indicates that levels of diversity in
subterranean groups there may be underestimated owing to conservative
fusiform morphology of burrowing animals.

Keywords: cryptic species, frog, fossorial, Geraldton sandplain

INTRODUCTION Biju and Bossuyt, 2003 of India. Forwards burrowing

Southwestern Australia is a biodiversity hotspot has presumably arisen only once in the Arenophyrne-
(Cincotta et al. 2000; Myers et al. 2000), which is the Myobatrachus lineage (Emerson 1976; Menzies and
centre of diversity and endemism for many kinds of Tyler 1977; Trueb and Cannatella 1982; Davies 1984;
plants and animals (Hopper et al. 1996; Hopper and Tyler 1994).
Gioia 2004), including several lineages of frogs (Roberts “Cryptic” species are good evolutionary species
and Watson 1993). Within this region, ancient that are not recognised as such owing to their
radiations of species have occurred within the two morphological similarity to one or more described
oldest families of frogs, the Myobatrachidae and forms (Donnellan et al. 1993). Taxa with conservative
Limnodynastidae (Frost et al. 2006). Within the morphologies are especially difficult to detect and
Myobatrachidae, one particular monophyletic lineage can only be elucidated with genetic techniques or
contains three related monotypic genera: Arenophryne large series of specimens to enable morphologists to
Tyler, 1976, Myobatrachus Gray, 1841 and Metacrinia tease out subtle but consistent differences among
Harrison, 1927 (Read et al. 2001). All are restricted to forms. Homoplastic traits (i.e., widely distributed
the southwest and have direct-developing young, traits within a lineage) may be the result of adaptive
fossorial habits and do not hop. Metacrinia nichollsi convergence of traits owing to a similar pattern of
crawls among the moist leaf-litter of the southern natural selection acting in similar environments.
forests, whereas Myobatrachus gouldii and A. rotunda Species with adaptations to swimming or burrowing
have evolved subterranean habits including the are especially likely to harbour cryptic species as
evolution of a more fusiform shape (small head, short external morphology becomes more streamlined to
limbs, flattened body) and burrow forwards through enable the animals to move more efficiently through
sand. Forwards burrowing in anurans (> 4000 species) a liquid medium (i.e., water or sand).
has evolved independently only a few times: for A recent molecular genetic study by Edwards (2007)
example, species in the microhylid genera Copiula has revealed significant genetic structuring within
Mehely, 1901 and Choerophryne Van Kampen, 1914 of Arenophryne . Here, we present a detailed
New Guinea, Hemisus marmoratus Peters, 1854 from morphological analysis of variation within A. rotunda
Africa, Rhinophrynus dorsalis Dumeril and Bibron, and describe as new a second species to the south of
1841 from Mexico and Nasikabatrqchus sahyadrensis the populations of A. rotunda from Shark Bay.
122 P. Doughty, D. Edwards

METHODS interaction term indicated sex differences were

Table 1 shows the morphological variables more pronounced in A. rotunda . Head length
measured with their definitions and abbreviations. showed significant main effects and interactions,
We also calculated the following ratios: HL/SVL, mostly owing to a larger size for A. rotunda
HL/HW, EN/IN, EN/IO, TibL/SVL, TarL/SVL and females. Arm width also showed complex, although
TarL/TibL. Genetic analyses (Edwards in press) subtle, interactions. No characters of the rear limbs
indicated a clear disjunction of lineages within the differed significantly between the two taxa.
range of A. rotunda occurring between Edel Land Colouration between the two taxa was also
and Coolamia Station (Figure 1). We selected noticeably different. Northern Shark Bay pop-
approximately 30 adult specimens from within each ulations are pale white with black flecks (tending to
of the regions identified by the genetic analyses of form paravertebral rows) and often some red
each taxon for our morphological comparison. flecking. In contrast, southern populations are a
Visual examination of the frequency distributions much darker brown (also with broad darker
of traits indicated no obvious violations of paravertebral rows and some red flecking) with
normality and heteroscedasticity. A 2-way ANOVA usually a dark transocular bar present.
of SVL was carried out with region (or “species”) Based on the morphological observations
and sex as factors. We tested whether there were presented above and the molecular genetic analysis
significant differences of morphological characters of Edwards (2007), we describe the southern taxon
with 2-way ANCOVA with species and sex as of Arenophyrne as a new species.
factors and body size (SVL) as the covariate. When
factors or interactions were significant, we present
summaries for each category separately in Table 2. TAXONOMY
All specimens from Western Australian Museum
(WAM prefix excluded from registration numbers). Amphibia

Family Myobatrachidae Schlegel 1850

RESULTS Genus Arenophryne Tyler 1976
Table 2 summarises the morphological differences
between Arenophyrne from the two regions. Two
main differences between the regions were Arenophryne xiphorhyncha sp. nov.
apparent. First, individuals from the southern
Southern Sandhill Frog
region had narrower heads, smaller eyes and
Figures 2 and 3
shorter distances for EN, IO and IN. These
characters are likely to be highly correlated with a Material examined
reduction in head size. Second, frogs from the
southern region had significantly larger hands. Holotype
Several characters displayed complex interactions Australia: Western Australia: WAM R67321. An
or were not significant. Females were larger in both adult female collected at Cooloomia Station, Western
taxa, with no difference in body size although the Australia (27°01'S, 114°19'E – 17 km at 240° angle from

Table 1 Characters measured with abbreviations and explanations.

Character Abbrev. Explanation of Measurement

Adults

Snout-vent length SVL From tip of snout to posterior tip of urostyle

Inter-limb length ILL From axilla to groin
Head length HL From tip of snout to posterior edge of midpoint of tympanic fold
Head width HW Width of head at midpoint of tympanic fold
Eye-naris distance EN From anterior corner of eye to posterior edge of naris
Interorbital span IO Distance between anterior corners of eyes
Internarial span IN Distance between inner edges of nares
Eye length EyeL Anterior to posterior corners
Hand length HandL Tip of 2nd finger to proximal edge of palmar tubercle
Arm width ArmW Maximum width of forearm
Tibia length TibL Measured with leg in natural resting position, from knee to tarsus
Tarsus length TarL Measured with leg in natural resting position, from proximal end of tarsus to
proximal edge of inner metatarsal tubercle
Foot length FootL From tip of 4th toe to proximal end of inner metatarsal tubercle
A new species of Arenophryne 123

Figure 1 Map of coastal Western Australia showing distribution of Arenophryne rotunda and A. xiphorhyncha sp.
nov.

Cooloomia homestead) by J. Rolfe, S. D. Hopper, P. J. limbs with unwebbed fingers and toes, palmar and
Fuller and K. Cashin on 19 September 1979. plantar surfaces bearing numerous tubercles,
tympanum absent, ground colour not yellow or
Paratypes pink, direct developing larvae and forward-
Australia: Western Australia: WAM R67320 and burrowing locomotion.
R67323 (females) details as for holotype; R123485 and Arenophryne xiphorhyncha is distinguished from
R126270 (males) 50 km N Kalbarri – Carnarvon Basin A. rotunda by narrower head with sharper canthal
survey site ZU5 (27°15'25"S, 114°11'21"E); R123554 region, smaller and less protruding eyes, larger
and R126251 (males) 50 km N Kalbarri – Carnarvon hands and darker colouration.
Basin survey site ZU4 (27°15'24"S, 114°9'11"E);
R126245 (male) 50 km N Kalbarri – Carnarvon Basin
Description
survey site ZU2 (27°15'41"S, 114°1'48"E); R126261
(female) 50 km N Kalbarri – Carnarvon Basin survey Holotype
site ZU1 (27°15'42"S, 114°1'9"E); R165815 (female) Measurements (mm): SVL – 30.0; ILL – 15.4; HL –
Sandy Junga Pits (27°49'59"S, 114°21'53"E); R165821 7.9; HW – 10.3; EN – 1.6; IO – 4.2; IN – 2.0; EyeL –
(female) 10 km NW of Murchison House Station 2.8; HandL – 6.1; ArmW – 2.8; TibL – 4.4; TarL – 3.8;
(27°36'22"S, 114°09'27"E). FootL – 7.1, HL/SVL – 0.26, HL/HW – 0.77, EN/IN –
0.80, EN/IO – 0.38, TibL/SVL – 0.14, TarL/SVL –
Diagnosis 0.12, TarL/TibL – 0.86.
A member of the genus Arenophryne based on Body dorsoventrally compressed and ovoid when
compressed ovoid body shape, small head with viewed dorsally (Figure 2A). Skin loose with
blunt snout with thickened epithelial tissue, short slightly raised bumps and folds scattered along
124 P. Doughty, D. Edwards

Table 2 Summaries of characters and ratios measured for Arenophryne xiphorhyncha and A. rotunda. Mean±S.D.
(range). N = 30 for both taxa unless noted. See Table 1 for abbreviations. SVL was tested with a 2-way
ANOVA. 2-way ANCOVAs (factors – species and sex, covariate – SVL) were carried out (see text for
explanation) and reported in the last column. Unless noted, sex and all species X sex interaction terms were
not significant with alpha = 0.05. Key: NS – not significant: P > 0.10, (*) 0.05 < P < 0.10, * P < 0.05, ** P < 0.01,
*** P < 0.001, **** P < 0.0001.

Character A. xiphorhyncha A. rotunda Statistics

N = 30 N = 30

SVL Female (N = 20): Female (N = 23): Spp.: F1,56 = 0.445NS

30.2±2.8 32.3±2.9 Sex: F1,56 = 14.811***
(26.0-36.0) (26.5-39.0) Spp. X Sex: F1,56 = 4.278*
Males (N = 10): Males (N = 7):
28.9±1.7 27.9±1.4
(26.5-32.0) (26.5-30.5)

ILL 12.5±2.1 13.8±2.0 Spp.: F1,54 = 2.704NS

(9.5-17.1) (9.2-19.7) SVL: F1,54 = 47.8****
N = 28

HL Female (N=20): Female (N=21): Spp.: F1,53 = 4.362*

8.0±0.4 8.7±0.6 Sex: F1,53 = 2.531(*)
(7.2-8.8) (7.4-9.6) Spp. X Sex: F1,53 = 5.167*
Males (N=10): Males (N=7): SVL: F1,53 = 35.35****
7.9±0.6 7.7±0.3
(6.9-8.7) (7.2-8.1)

HW 9.6±0.6 10.5±0.6 Spp.: F1,54 = 44.7****

(8.6-10.9) (9.1-11.8) SVL: F1,54 = 30.3****
N = 28

EN 1.6±0.2 1.9±0.2 Spp.: F1,56 = 30.5****

(1.4-2.0) (1.7-2.4) SVL: F1,56 = 6.90*

IO 4.2±0.2 5.0±0.3 Spp.: F1,56 = 207.8****

(3.8-4.8) (4.5-5.7) SVL: F1,56 = 42.9****

IN 2.0±0.2 2.2±0.2 Spp.: F1,56 = 27.8****

(1.7-2.2) (1.8-2.9) SVL: F1,56 = 16.6****

EyeL 2.9±0.2 3.4±0.3 Spp.: F1,56 = 37.2****

(2.4-3.4) (2.8-4.2) SVL: F1,56 = 16.1***

ArmL Female (N = 20): Female (N = 23): Spp.: F1,55 = 0.533 NS

3.1±0.4 3.0±0.3 Sex: F1,55 = 8.141**
(2.4-4.0) (2.5-3.7) Spp. X Sex: F1,55 = 5.167*
Males (N = 10): Males (N = 7): SVL: F1,55 = 18.85****
2.5±0.3 2.6±0.3
(2.0-2.8) (2.0-2.9)

HandL 5.9±0.4 5.6±0.4 Spp.: F1,55 = 18.3****

(5.0-6.8) (4.8-6.5) SVL: F1,55 = 7.36**
N =29

TibL 5.6±0.6 5.6±0.7 Spp.: F1,49 = 0.358NS

(4.4-7.0) (4.4-7.2) SVL: F1,49 = 11.179**
N = 28 N = 25

TarL 3.9±0.5 3.8±0.5 Spp.: F1,49 = 0.103NS

(3.0-5.0) (2.7-4.6) SVL: F1,49 = 2.008NS
N = 28 N = 25

FootL 7.2±0.5 7.4±0.6 Spp.: F1,45 = 0.033NS

(6.1-8.4) (6.3-8.4) SVL: F1,45 = 11.015**
N = 27 N = 22
A new species of Arenophryne 125

Table 2 (cont.)

Character A. xiphorhyncha A. rotunda Statistics

N = 30 N = 30

HL/SVL 0.27±0.02 0.27±0.02

(0.23-0.29) (0.24-0.30)
N = 28

HL/HW 0.83±0.03 0.80±0.04

(0.77-0.90) (0.73-0.87)
N = 28

EN/IN 0.83±0.07 0.85±0.10

(0.71-0.95) (0.66-1.20)

EN/IO 0.39±0.04 0.38±0.03

(0.33-0.40) (0.33-0.46)

TibL/SVL 0.19±0.02 0.18±0.02

(0.14-0.24) (0.14-0.22)
N = 28 N = 25

TarL/SVL 0.13±0.02 0.13±0.02

(0.10-0.17) (0.09-0.17)
N = 28 N = 25

TarL/TL 0.70±0.08 0.69±0.09

(0.66-0.88) (0.52-0.85)
N = 28 N = 25

body and tending to form vertebral, paravertebral Colouration in preservative

and dorsolateral ridges; rugose along lateral Light brown dorsum with darker dorsolateral
surfaces. stripes. Dark grey patches on shoulders. Top of
Head small. Snout narrow with moderate canthus head pale with almost white snout. Limbs cream
rostralis (Figure 3C). Thickened epithelial stratum with pale digits. Belly pale brown with lighter
corneum covers the end of the snout. Eyes not cream colour towards flanks; chin cream
projecting far beyond outline of head or body. (unpigmented).
Nostrils near end of snout and directed upwards.
Mouth wide, terminating below eyes. Vomerine Variation
teeth absent. Tongue narrow and long. No visible Females were larger than males (Table 2), as is the
tympanum, but with distinct tympanic fold case for most anurans. A lack of smaller size classes
posterior to eye. Raised skin with glandular precluded an analysis of size at maturity. Overall,
appearance between mouth and forelimbs (Figure there was little variation in head or limb
3A). Urostyle not projecting; cloaca directed proportions among individuals, including no
posteriorly and downwards. pronounced sex-based differences in shape. More
Limbs massive and extremely short. Forelimbs pronounced body shape differences among the
stout and covered in loose skin with elbow barely preserved specimens was due to the fullness of the
discernible. Hands robust with first three fingers gut. Many individuals had guts filled exclusively
extremely thickened and numerous tubercles with ants which gave them a plump appearance.
scattered on palm including large palmar tubercle; Individuals varied in the rugosity of the skin from
4th finger extremely reduced (Figure 3F). Fingers nearly smooth to raised folds of skin tending to
with strong fringes, including ridges formed on the form ridges along the dorsolateral stripes (as in the
sides of the hand along the palm and inner edge of holotype). Likewise, ventral surfaces ranged from
1st finger and outer edge of 4th finger. Finger length: almost smooth to moderately granular, although
2>3>1>4. Legs also stout and thick and covered with this appeared to vary with the preparation and age
loose-fitting skin. Feet robust with strong fringes of specimens.
and numerous tubercles on the plantar surface
(Figure 3G). Toe length: 4>3>5>2>1 (1 st toe Colour in life
extremely reduced to almost the size of the plantar Live A. xiphorhyncha (Figure 2A) have a medium
tubercle). to dark brown dorsum with pale limbs, flanks and
126 P. Doughty, D. Edwards

A B

Figure 2 A, Arenophryne xiphorhyncha sp. nov. from type location (Coolomia Station, Western Australia);
B, A. rotunda also from the type location (False Well Entrance, Shark Bay, Western Australia).
Photographs by Brad Maryan.

snout. There are usually conspicuous darker the center of the belly and faded distally towards
paravertebral stripes on the back, along with dark the head, flanks and legs.
irregular markings. There is a lighter vertebral area
with a thin clearly demarcated yellow to cream Etymology
stripe running from the back of the head to the The specific name is a Latinized version of the
urostyle where it is more clearly seen. Raised Greek xiphos (sword) + rhynchos (nose or snout) in
tubercles and folds on the dorsum are often tipped reference to the sharper snout of A. xiphorhyncha
with the pale ground colour. There are often compared to A. rotunda. For common names, we
scattered deep red flecks present on the dorsum and suggest for A. xiphorhyncha the “southern sandhill
some yellow flecks present on the sides, especially frog” and for A. rotunda the “northern sandhill
near the groin. The belly is pale with stippling or frog”.
irregular blotching with a semi-translucent
abdomen. Comparisons with other species
Arenophryne xiphorhyncha occurs near four
Colour in preservative other myobatrachid frogs on the central western
Pale to dark brown with irregular light and dark coast. Although the call of A. xiphorhyncha is not
flecking and darker vertebral and paravertebral known, many myobatrachids have very similar calls
longitudinal stripes or bands discernible. Head consisting of a short harsh rasp, including A.
slightly paler than body colour with very pale rotunda , Myobatrachus gouldii , Metacrinia
snout. Thin yellowy vertebral stripe visible on nichollsi, all Pseudophyrne Fitzinger, 1841 and
posterior half of dorsum. Red flecks present only many Uperoleia Gray, 1841 (Roberts 1984). We
on recently-preserved (< 2 y) specimens. Para- anticipate that A. xiphorhyncha will have a similar
vertebral stripes beige to light-brown to blue-grey, call, although this remains to be documented.
from faintly expressed to very dark and Pseudophryne guentheri is known from the area
conspicuous. Vertebral area between paravertebral and can be distinguished from A. xiphorhyncha by
stripes ranges from pale background colour to the following traits: body much less stout, snout
nearly the same darkness as the paravertebral and eyes more prominent, longer and more slender
stripes in some specimens. Canthal stripe passes limbs with long fingers and toes, large metatarsal
through eye to continue as dorsolateral stripe. Side tubercles, often large dark blotches on back,
of head below canthal stripe and eye has the pale backwards burrowing, lays eggs. Pseudophyrne
ground colour. Dorsolateral stripe ranges from a occidentalis occurs just to the north and east of the
thin line angled downwards from shoulder to groin known distribution of A. xiphorhyncha. In addition
with faint stippling below to nearly a solid dark to the characters listed for P. guentheri , P.
stripe along flank. Forelimbs and hands pale. Rear occidentalis also has an orange patch on the snout
thigh and tibia same as dorsal colouration, but with between the eyes and usually on the elbows and
pale feet (as for forelimbs and hands). Belly patterns rump.
were variably stippled with black, but in general Myobatrachus gouldii is known to occur just to
the pattern was for a pale background upon which the south (Eradu) of A. xiphorhyncha . It is
was darker stippling ranging from diffuse to distinguished by its pink to yellow skin, more
markedly blotched. Stippling was concentrated in reduced and fusiform head and attains a larger
A new species of Arenophryne 127

Figure 3 Diagrams of the lateral view of the head (A), anterior view (B), dorsal view (C), hands (F) and feet (G) of the
holotype (WAM R67321) of Arenophyrne xiphorhyncha sp. nov. Anterior (D) and dorsal (E) views of the
head of A. rotunda are provided for comparison (WAM R68348).
128 P. Doughty, D. Edwards

body size (50 mm vs. 30 mm SVL in A. xipho- and subtle morphological differences between the
rhyncha). species indicates that there could be other cryptic
Arenophyrne xiphorhyncha differs from its species of frogs and reptiles that show reduction of
congener A. rotunda by, in general, possessing morphological traits owing to adaptations for
traits indicating a stronger commitment or history burrowing in sand. For example, species in the
of burrowing habits. The head of A. xiphorhyncha gekkonid genus Aprasia are subterranean
is more streamlined including smaller and less burrowers that inhabit the coastal sands in
protruding eyes, shorter distances between the eyes southwestern Australia up to the Pilbara region.
and nostrils, and a sharper snout. The hands of A. Recent morphological and genetic research has
xiphorhyncha are larger, have more tubercules and revealed many cryptic species within these forms
the ridges on the sides of the hand are more (B. Maryan, K. Aplin and M. Adams, unpublished
developed than those of A. rotunda . The data). Western heath dragons (genus Rankinia
colouration of A. xiphorhyncha is much darker than Wells and Wellington, 1985) also exhibit an affinity
that of A. rotunda (Figure 2). to isolated sandy habitats in the southwest
(including the unique ability within Australian
Habitats, feeding and breeding biology dragons to “shimmy-bury” in the sand – Greer
Arenophryne xiphorhyncha inhabits sandy 1989). Western heath dragon populations showing
regions within the Geraldton sandplain. Frogs deep historical divergences genetically (Melville
presumably shelter underground during the day, and Doughty in press). However, the sister group
and emerge at night to feed on the surface at to Arenophryne – the obligate sand-dwelling
suitable times of the year (autumn – spring). Obser- forwards-burrowing turtle frogs (Myobatrachus
vations of gut contents of preserved specimens gouldii) – are distributed over a much wider area in
were entirely of ants, but more detailed analyses southwestern Australia, but do not exhibit
may yield a wider range of prey types. significant breaks in phylogeographic structure over
Nothing specific is known of the breeding biology their range (S. Keogh, P. Doughty, M. Adams and
of A. xiphorhyncha, although the breeding biology D. Edwards, unpublished data).
of A. rotunda is moderately well-known (Roberts Climate induced sea level fluctuations during the
1984; Anstis et al. 2007). In A. rotunda, males call in Plio-Pleistocene, resulting in coastal dune evolution
late winter and spring in response to rain. Pairs in the region (Hocking et al. 1987), have been
form and spend the summer together when frogs hypothesised as drivers of speciation within
are inactive owing to high temperatures and lack of herpetofauna of the Shark Bay and wider Carnar-
rain. Eggs are deposited in autumn in about 80 cm von Basin region (Storr and Harold 1980; Rabosky
of sand and hatch over two months later. et al. 2004). Fluctuating sea levels are plausible
All previous reports of the biology of explanations for vicariance in species with disjunct
Arenophryne have been on A. rotunda from Shark populations across the northern Carnarvon Basin,
Bay. Presumably, many of the habits, ecology and such as Rankinia (Melville and Doughty, in press)
breeding biology of A. xiphorhyncha will be similar and several other skink and gecko species (Storr and
to A. rotunda, but further studies need to be con- Harold 1978, 1980). However, divergence estimates
ducted to test this supposition. dating the split within Arenophryne (Edwards
2007) suggest that speciation predates the Plio-
Pleistocene sea level fluctuations. Molecular clock
Distribution estimates can be subject to error (Rambaut and
Arenophryne xiphorhyncha is only known from a Bromham 1998), however, the above date provides
broad strip of sandplain north of Geraldton and an estimate that is correlated with known climatic
south of Shark Bay, Western Australia (Figure 1). and geological changes.
The distance between the northern and southern- The formation of the Victoria Plateau, in
most locality records is approximately 120 km. It is combination with sweeping aridity, is likely to
worth noting that the description of the southern have led to the Late Miocene divergence between
Arenophryne as a separate species reduces the the two Arenophryne species (Edwards 2007).
range of true A. rotunda considerably. During the late Miocene, tectonic instability
Arenophryne rotunda is now confined to the white resulted in the reactivation of pre-existing faults
coastal dunes from the northern tip of Dirk Hartog and the uplift and formation of the Victoria
Island to near the base of the Edel Land peninsula – Plateau, with the Victoria Plateau uplifted by as
approximately 150 km. much as 60m in the Kalbarri region (Haig and
Mory 2003). The northern border of the Victoria
Plateau corresponds to the geographic position of
DISCUSSION the genetic break between the two Arenophryne
The detection of a second species within species. The thick coastal sand deposits of the Edel
Arenophyrne based on molecular genetic results group (common in the area today) were not
A new species of Arenophryne 129

formed until the Plio-Pleistocene (Hocking et al. less time spent on the surface, and hence less need
1987), therefore an alternative avenue for dispersal for the ground colour to match the substrate
would not have been available for obligate sand- background to avoid predation by visually-oriented
dwelling forms. predators.
The current distribution of A. rotunda is restricted The evolutionary precursor to forwards-
to the coastal white sand dunes along the Edel and burrowing through sand may have been the
Shark Bay Peninsulas, while the distribution of A. crawling habits of the sister taxon to all three
xiphorhyncha covers much more variable in soil burrowing forms, Metacrinia nichollsi. This species
types (Figure 1). South of the Murchison Gorge, is similar to Pseudophryne in its ground-dwelling
populations occur on black sandplain. Immediately habits, including walking or crawling instead of
north of the Murchison Gorge and up to the hopping. Metacrina lives in deep leaf litter in
Zuytdorp coastal region, populations occur on southwest Australia, and diving in to this substrate
yellow sandplain. Further inland of the Zuytdorp may have led to the development of forwards-
coast and up into the Cooloomia region, burrowing in Arenophryne and Myobatrachus. The
populations occur on siliceous red sandplain and forwards-burrowing asterophryine and spheno-
dune systems. The morphological differences phrynine microhylids of Papua New Guinea also
between A. rotunda and A. xiphorhyncha may be have close relatives that occur in dense leaf litter,
representative of a history tied to coastal sands (in supporting this supposition (Menzies and Tyler
the case of A. rotunda) as opposed to a history tied 1977; Davies 1984). More comparative work on the
to sandplain complexes (in the case of A. evolution of forwards-burrowing habits and the
xiphorhyncha). attendant morphological adaptations such as
When considering the morphology and reduced head size, reinforced pectoral girdle,
appearance of A. xiphorhyncha with its close increase in arm and hand size, rotation of the angle
relatives, it appears to be intermediate between A. of the limbs for burrowing and other characters
rotunda and M. gouldii in its commitment to would be a fruitful area of study, especially given
burrowing habits. Although A. rotunda is a fully its multiple origins within the anurans (Emerson
subterranean species like the other two, it retains a 1976; Menzies and Tyler 1977; Trueb and Canatella
more globular “frog-like” appearance. In contrast, 1982).
A. xiphorhyncha has a smaller head with smaller
less protruding eyes, shorter distances between all
facial distances (Table 2) and a sharper canthus, all ACKNOWLEDGEMENTS
of which present a smaller surface area when We thank Claire Stevenson for the drawings and
pushing forwards through the sand. Morphological map, Brad Maryan for providing the photographs,
differences between the two Arenophryne species Alicia Carlisle for help in the laboratory, and Dale
may have evolved in response to the relative Roberts, Glenn Shea and Mike Tyler for helpful
difficulty of pushing through the heavier yellow discussions.
and red calcareous sands of the Geraldton
sandplain in the case of A. xiphorhyncha compared
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A new species of Arenophryne 131

APPENDIX
Comparative material examined. All specimens from the Western Australian Museum, Welshpool (R prefix
omitted below).

Arenophryne xiphorhyncha
Females – 66444, 121780, 123495, 123497, 123499, 123500, 126244, 126246, 126254, 126259, 126271, 126272,
126288, 165822.
Males – 123550, 123560, 126262, 126267, 126278.

Arenophyrne rotunda
Females – 55206, 68350, 114066, 114083, 114084, 122520, 126156, 126158, 126159, 146480, 157824, 157825,
157826, 157828, 157831, 157832, 157833, 157834, 165796, 165804, 165805, 165806, 165809.
Males – 68348, 68349, 87852, 87853, 165803, 165808, 165810.