Taxonomy of Rock Wallabies, Petrogale (Marsupialia : Macropodidae). I. A Revision of the Eastern Petrogale with the Description of Three New Species
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Aust. J. Zool., 1992, 40, 605-25
Taxonomy of Rock Wallabies,
Petrogale (Marsupialia :Macropodidae). I.
A Revision of the Eastern Petrogale
with the Description of Three New Species
M. D. B. Eldridge and R. L. closeA
School of Biological Sciences, Macquarie University,
N.S.W. 2109, Australia.
A Current address: School of Business and Technology,
Univesity of Western Sydney, Macarthur, P.O. Box 555,
Campbelltown, N.S.W. 2560, Australia.
Abstract
The taxonomy of Petrogale has been in a state of flux for many years. The eight chromosome races
of the eastern Petrogale radiation are currently placed in four species. However, several of these 'species'
contain chromosomally unrelated taxa. In this paper a species definition for Petrogale is proposed that
allows for some gene flow between species but requires a species to maintain a substantial and distinct
genetic identity. When this definition was applied to the eastern Petrogale eight 'good' species were
identified. Thus we now consider the eastern Petrogale complex to consist of P. penicillata, P, herberti
(formerly P. penicillata herberti), P. inornata, P. assimilis, P. sharmani, sp. nov. (formerly the
Mt Claro race), P. mareeba, sp. nov. (formerly the Mareeba race), P. godmani and P. coenensis,
sp, nov. (formerly the Cape York race). Several of these taxa are cryptic species and the primary means
of identification used was chromosome number and morphology. However, genic data were useful in
establishing whether each taxon should be regarded as a separate species.
Introduction
The indigenous Australian genus Petrogale (rock wallabies) includes small (0.9 kg) to
medium-sized (9 kg) macropods,,that are found throughout Australia and on some offshore
islands (Strahan 1983). As their name implies, rock wallabies live in rocky habitats,
preferring steep rocky slopes, cliffs, gorges, rocky outcrops and boulder piles (Sharman and
Maynes 1983a). Considerable variation in size, pelage characteristics and skull morphology
has led to the description of 23 Petrogale taxa in the last 160 years (Briscoe et al. 1982).
In that time there has been little agreement as to the status of these taxa: Iredale and
Troughton (1934) recognised 11 species, Marlow (1965) nine, Ride (1970) eight and Poole
(1979) seven.
More recently, a multifaceted study of Petrogale was begun by Professor Geoff Sharman
and his colleagues at Macquarie University. During an Australia-wide survey, members of
the Macquarie group attempted to re-collect all described Petrogale taxa at or near their type
localities. On the assumption that the karyotypes of newly collected specimens were the same
as those of the type specimens, the karyotype attributable to each taxon was identified.
Additional specimens from other localities were thus referred to a particular taxon on the
basis of sharing a similar karyotype to that found at the relevant type locality (Sharman
et al. 1990).
These studies have indicated that only some Petrogale taxa can be readily distinguished
by traditional mammalian taxonomic characters (i.e. pelage and skull characteristics).
For example, morphologically distinct populations of Petrogale from northern Australia,
previously described as five separate taxa (P. brachyotis brachyotis, P. brachyotis signata,M. D. B. Eldridge and R. L. Close
P. longrnani, P. venustula and P. wilkinsi), were all found to be chromosomally similar and
so were placed in a single species, P. brachyotis (Sharman and Maynes 1983~;Sharman
et al. 1990). Also, variation in skull morphometrics within P. assirnilis assirnilis, P. godrnani
godrnani and the Mareeba, Mt Claro, and Cape York races of north-eastern Australia is
considerable: while skull size and morphology tend to be uniform within a particular colony,
they can vary significantly between colonies within each taxon. As the variation within taxa
is often greater than that found between taxa, it is difficult to use these characters to
distinguish between taxa (R. L. Close and D. Haig, unpublished data). Similarly, these taxa
can not be reliably distinguished by pelage characteristics (Sharman and Maynes 1983b;
Maynes and Sharman 1983). However, each of these five taxa is clearly distinct and readily
identified by characteristic chromosome (Sharman et al. 1990) and genic (Bee and Close
1992) markers.
Thus these genetic techniques have proved useful in identifying distinct taxa and in deter-
mining the geographic location of taxonomic boundaries between parapatric populations.
The most definitive of these techniques, in terms of its ability to readily and reliably identify
Petrogale taxa, has been cytogenetics. This initially involved identifying differences in
chromosome number, size and morphology (i.e. the position of the centromere) using con-
ventionally stained preparations (Briscoe et al. 1982; Sharman et al. 1990). More recently,
G-banding, a technique that allows homologous chromosomes or chromosome arms to be
identified by their characteristic banding patterns, has been used to confirm the identity
of the chromosomes involved in rearrangements and to detect cryptic and/or internal
chromosome changes (Eldridge et al. 1988, 1989, 1990, 1991a, 1992a, 19926).
Phylogenies constructed from the G-banding data (Eldridge et al. 1991b) have revealed
the inadequacy of present taxonomy. For example, P. g. godrnani and the Cape York
race are currently placed in a single species, P. godrnani (Sharman and Maynes 1983b);
however, the G-banding data show that these taxa are not closely related (Eldridge et al.
1991b) and are sufficiently distinct from each other to be regarded as separate species
(Eldridge et al. 1989). Similarly, the chromosomes of the Mareeba and Mt Claro races are
sufficiently distinct from P. a. assirnilis (Eldridge et al. 1991b) to warrant their removal
from P. assirnilis (Eldridge et al. 1989).
These easily identified inadequacies are most likely symptomatic of the need for a general
revision of the genus and particularly the eastern taxa. T o properly evaluate the taxonomic
status of each Petrogale taxon, the chromosome phylogeny will be compared with the
available genic data [i.e. electrophoretic variation (Table I), mitochondria1 DNA (mtDNA)
restriction morphs (Bee and Close 1992) and hybrid breeding trials (Table 2)]. These com-
parisons should result in a set of criteria by which Petrogale species can be readily identified.
As the taxonomy of Petrogale has been in a state of flux for many years some clarification
of species-level taxonomy should greatly assist the researcher and naturalist, as well as
wildlife- and conservation-authorities.
A Species Definition for Petrogale
The widely held biological species concept (Dobzanhsky 1937; Mayr 1940) emphasises the
need for gene flow between populations within a species but requires reproductive isolation
between species. Amongst the eastern Petrogale taxa, studies of contact zones and the fertility
of hybrids produced in captivity have shown that reproductive isolation amongst many taxa
is incomplete (Table 2). Thus, in terms of a strict interpretation of the biological species
concept (e.g. Key 1981), most taxa in the eastern Petrogale radiation could be placed in a
single species. This would be inappropriate given that many of these taxa are just as well
differentiated as other species in which no introgression has been detected.
It is often regarded as a tacit assumption that interbreeding populations must either be
conspecific or at most nearest relatives (Wiley 1981), but in Petrogale introgression has
frequently occurred between taxa that are not nearest relatives. Rosen (1979) has pointedTable 1. Diagnostic electrophoretic loci for the eastern Petrogale taxa
For each taxon the characteristic allele(s) at each locus idare given, in bold, on the first line. Minor alleles are indicated on the second line. lntrogressedA alleles
are shown in parentheses. S, slow; N, normal; F, fast; vF, very fast. Where more than four alleles were present at a locus they were numbered consecutively,
with 1 being the most cathodal allelic form. Additionally, all eastern Petrogale taxa were characterised by the N allele at the Sdh, Ada-C, or-Gpd, Sod-A, Sod-C,
Ca, Mdh-A, Mdh-C, Gpd, Ldh-B, Pgm-C, Tf, Pgk, Fum, Ak-A, Ak-C, Idh-C, Me-C, Pep-A, Pep-b, AAT-C, Gpt-mus, Pk-mu, Acp-A, Enol-A, Enol-C and
Gdh loci. See Appendix for details of locus abbreviations. (Data from D. A. Briscoe.)
Taxon Presumptive gene loci
Pgi Ada-A 6Pgd Ldh-A Pgm-A Acon-A Acon-C or-Gal Idh-A Mpi Me-A PepD Alb Cat AAT-F AAT-2 Gpt Gda Pk-liv Acp-C Xdh
Cape York race N 2 N N N N N S N 5 N N N S / N N N N F N S / N N
P. g. godmani N 2 SIN N N N N S N 5 N N N S / N N N/F N/FF N N N/F
F (3) F IF) /N) 2 /F) IS) /FJ (8 /N) /S)/F S
Mareeba race N/F 3 N N N N N/F N N 5 N/F N/F S/N N/F N S/N/FN N N/F S/N N
12) /WF /F) 6) /F)
Mt Claro race N 3 S/N/F S / N N N N N N/F 5 N S/N N N N S/N/F N N N N N/F
/F) F S
P. a. assirnilis N/F 3 N/F N N N N/F N N 5 N N N N N S/N/FN N N N N/F
25 S S/F 2 /s)/F F S//F) S/F S
P. inornata S/N/F3 N N N N N N N 5 N N N N N N / F N F N N N
2 F F F F F
P.p.herberti N 3 N N N N N N N 2 N N/F N N S/N N N N N N N
(F) 1 S//F) F F /@ F (3)/5 F F S/F
P.p.peniciNata F/vF 3 N N S/N F F N N/F 3 N N N N S/N/F N/F N N N N N
/N) 1 F /N) (2)/3 S
A Where an allele characteristic of one taxon was found in an adjacent taxon, it was assumed to have introgressed only when the allele was present at or near the
contact/hybrid zone and was not found elsewhere in the taxon.M. D. B. Eldridge and R. L. Close
Table 2. Summary of data on the fertility of Petrogale hybrids and the genetic divergence of
their parental taxa
FGD, Tied gene difference (from Table 1); * indicates that evidence of introgression has been
found at the contact zone of these taxa (Table 1; Bee and Close 1992). Data on hybrids from
Sharman et al. (1990) and R. L. Close (unpublished)
Parental taxa Chromosome FGD Fertility of hybrids
heterozygosity Male Female
Cape York unknown unknown *
godrnani
godrnani sterile subfertile *
Mareeba
Mareeba sterile unknown a
Mt Claro
Mt Claro sterile unknown *
assirnilis
Mareeba sterile subfertile
assirnilis
assirnilis sterile unknown *
inornata
inornata unknown unknown *
herberti
herberti unknown fertile or *
penicillata subfertile
herberti sterile unknown
Mareeba
assirnilis sterile subfertile
penicillata
Mareeba sterile unknown
penicillata
persephone unknown fertile or
xanthopus subfertile
godrnani sterile unknown
purpureicollis
out that the ability to interbreed may be due to the retention of plesiomorphic characters
and therefore can not be considered as prima facie evidence for conspecificity or sister-
species relationship. Wiley (1981) regards hybridisation and introgression between unrelated
populations (which must represent secondary contact) as 'of no relevance to a decision as
to their species status'. However, the characteristics of hybrid/contact zones between nearest
relatives (whether they be primary or secondary) can be useful, when making decisions as
to specific status.
Therefore, introgression in Petrogale may not simply represent evidence of recent
speciation events but may also result from taxa retaining the ability to interbreed. Table 2
strongly suggests that wherever Petrogale populations become parapatric some hybridisation
will occur and that some of the resultant female hybrids will be at least partially fertile.
Many distinct macropod species are able to hybridise (Close and Lowry 1990). The
recently reported potential for five of these interspecific macropod hybrids to be fertile
(Close and Lowry 1990) is consistent with the notion that introgression can take place
between good species. Introgression does occur across taxon boundaries in eastern PetrogaleTaxonomy of Rock Wallabies. I.
but it varies in its extent (Briscoe et al. 1982; Bee and Close 1992). It is difficult, then, to
decide arbitrarily the point at which gene flow between two populations becomes sufficiently
reduced to warrant their being given specific status.
In Petrogale, a species is probably best defined as the taxon that exhibits sufficient
reproductive isolation from other taxa to maintain a substantial and distinct genetic identity,
even in the presence of some introgression from other (parapatric) taxa. Thus, at a contact
zone between presumptive species of Petrogale, if the hybrid zone is narrow and intro-
gression limited, then the two taxa are effectively maintaining separate genetic identities
and the researcher is justified in recognising two species. If, however, two taxa tend to merge
into each other and hybridise over a wide zone then they would most likely represent two
geographical variants of the same species.
A Revision of the Eastern Petrogale
The largest group of Petrogale is the widespread and diverse lateralis/penicillata group
(Briscoe et al. 1982). Within this group, two main complexes can be differentiated: a western
complex, comprising the races of P. lateralis, and an eastern complex distributed in the
ranges of the east coast of Australia (Briscoe et al. 1982; Eldridge et al. 1991b). The eight
taxa of the eastern complex form a virtually unbroken chain of parapatric taxa (Barker and
Close 1990) from Cape York, in the north, to Victoria, in the south (Eldridge et al. 1991b).
Although these taxa are amongst the most intensively studied Petrogale, their inter-relation-
ships remain confusing, with much of the genetic data being contradictory.
Currently, four species of Petrogale are recognised amongst the taxa of the eastern
radiation: P. godmani (containing, on geographical criteria, P. g. godmani and the Cape
York race), P. assimilis (containing, on geographical criteria, P. a. assimilis and the Mareeba
and Mt Claro races), P. inornata, and P. penicillata (containing P. p. penicillata and
P. p. herberti) (Calaby and Richardson 1988).
The species definition we have proposed for Petrogale is most readily assessed in para-
patric or syrnpatric taxa, since this geographical arrangement gives taxa the potential to
hybridise and therefore exchange genetic material. In parapatric taxa it is possible to assess
the extent of reproductive isolation between populations by examining the distribution of
characteristic genetic markers at the contact/hybrid zone between them. Thus the extent of
gene flow between the populations can be estimated and a decision made as to whether the
taxa are maintaining their respective genetic identities.
A comparison of the chromosomal (Briscoe et al. 1982; Sharman et al. 1990), electro-
phoretic (Table 1) and mtDNA (Bee and Close 1992) data shows that each race in the
eastern Petrogale radiation is identifiable by characteristic chromosomal and genic markers.
Additionally, at all examined contact zones, some leakage of genetic markers across taxon
boundaries is apparent; however, in all cases these contact zones were narrow and typically
the introgressed markers are found only in animals located at the contact/hybrid zones
(Table 1; Briscoe eta!. 1982; Bee and Close 1992). Additionally, many characteristic markers
were not found outside the taxon boundaries. Thus, each race of the eastern Petrogale
radiation can be seen to be maintaining a separate genetic identity and therefore should be
regarded as a distinct species.
It is useful to be able to reinforce species decisions based on genetic data by conducting
breeding trials to examine the fertility of the hybrids produced between presumptive species.
If these hybrids prove to be non-viable, or viable but sterile, or partly sterile, then the
hypothesis that two species are represented is reinforced. Any other result is essentially
equivocal. If no hybrids are produced then either the two taxa are not reproductively
compatible or they may just be difficult to breed in captivity. The production of fully
fertile hybrids does not prove that the taxa involved are conspecific, since reproductive
isolation may have been broken down by the artificial conditions, and under natural
conditions the taxa would never interbreed.M. D. B. Eldridge and R. L. Close
The data that have been obtained on the fertility of Petrogale hybrids are then particu-
larly useful. The extent of genetic divergence and the fertility of all known Petrogale
hybrids, both from naturally hybridising populations and from breeding trials in captivity,
are shown in Table 2. The results clearly follow Haldane's rule in that male hybrids are
more severely affected than female hybrids; in fact, no fertile male hybrid has been found.
Moreover, some females in most hybrid classes were at least subfertile (Table 2). The
ability of some female hybrids to be occasionally fertile and produce backcross offspring is
remarkable (given the chromosomal and/or genic differences that separate some taxa),
but is consistent with the finding that introgression has occurred across most contact zones
(Table 2).
Where no breeding data for specific pairs of taxa are available, it is frequently possible
to infer the extent to which the genic and chromosome differences present in the taxa of
interest will affect fertility in their hybrids by correlating the effect that similar genetic
differences have on the fertility of hybrids between other taxa for which data are available.
The G-banding data show that the Cape York race and P. g. godmani are not closely
related (Eldridge et al. 1989); however, the electrophoretic and mtDNA data are not
consistent with this (Table 1; Bee and Close 1992). Regardless of this contradiction, the
extent of chromosomal differences between the Cape York race and P. g. godmani suggests
that hybrids would have reduced fertility (Table 2), with male hybrids being sterile and
female hybrids most likely being subfertile. As these taxa represent two independent chromo-
somal lineages (Eldridge et al. 1991b), and gene flow between them would be restricted by
their chromosomal differences, they should be regarded as separate species (despite their
genic similarity) (Table 1). Similarly, Baverstock et al. (1986) regarded Rattus colletti and
Rattus villosissimus as separate species because of reduced fertility of hybrids (Baverstock
et al. 1983), despite the fact that they found no fixed gene differences between them.
Despite having similar mtDNA (Bee and Close 1992), P. g. godmani and the Mareeba
race clearly represent distinct species, having major chromosomal (Eldridge et al. 1991a)
and electrophoretic differences (Table 1). Although these genetic differences indicate that
these two taxa are not nearest relatives, a narrow hybrid zone has been located (Briscoe
et al. 1982). Typically, male hybrids were sterile and female hybrids subfertile (Table 2).
Despite this barrier to gene flow, some introgression of characteristic genetic markers has
been detected (Table 1). Atypical markers, however, are found only in animals from the
hybrid/contact zone and both taxa are maintaining their respective genetic identities. These
taxa demonstrate the potential for introgression to occur between Petrogale taxa that are
not nearest relatives and are otherwise well differentiated.
The decision, based on G-banding data (Eldridge et al. 1989), to remove the Mareeba and
Mt Claro races from P. assimilis is consistent with the reported sterility of Mareeba race x
P. a. assimilis and Mt Claro racexP. a. assimilis males, the sterility or subfertility of
similar hybrid females (Table 2) and the lack of significant introgression across their
respective contact zones (Table 1; Bee and Close 1992).
The G-banding data indicate that the Mareeba and Mt Claro races are nearest relatives
(Eldridge et al. 1991b). An analysis of allozyme variation at their contact zone showed
evidence of introgression at only two loci (Acon-C, Idh-A) (Table 1). Moreoever, no
evidence of introgression was found for three other characteristic Mareeba-race alleles
(Pgi allele F, Me-A allele S , Cat allele F ) or for four characteristic Mt Claro alleles (Ldh-A
allele S, 6Gpd allele S , Pep-D allele S , Xdh allele F ) (Table 1). Additionally, an analysis of
mtDNA restriction morphs showed no evidence of introgression (Bee and Close 1992).
Thus these two taxa appear to be maintaining separate genetic identities and should therefore
be regarded as separate species. Additionally, a hybrid Mareeba race x M t Claro race male
was sterile (Table 2).
At the P. inornata/P. a. assimilis contact zone only minor leakage of genetic markers
has been detected (Table 1; Bee and Close 1992). It can thereforfe be concluded that these
two taxa are sufficiently reproductively isolated to be regarded as distinct species. Moreover,
a P. a. assimilisxP. inornata hybrid male was sterile (Table 2).Taxonomy of Rock Wallabies. I.
The chromosome data indicate that P. inornata and P. p. herberti are not closely related
(Eldridge et al. 1991b). The electrophoretic (Table 1) and mtDNA (Bee and Close 1992)
data reveal little evidence of gene flow between these taxa, although P. inornata and
P. p. herberti are clearly in physical contact (Barker and Close 1990). The genetic differences
between these two taxa are greater than those between P. inornata and P. a. assimilis
(Table 2) and therefore could be expected to significantly reduce the fertility of hybrids.
There can be little doubt that P. inornata and P. p. herberti represent distinct species.
The mtDNA data (Bee and Close 1992) indicate that P. p. penicillata and P. p. herberti
are nearest relatives and they are currently regarded as conspecific (Calaby and Richardson
1988). However, they do not appear closely related electrophoretically (Table 1) and the
chromosome data is equivocal (Eldridge et al. 1991b). A hybrid zone between these taxa has
been found in south-eastern Queensland but only limited data exist as to the fertility of
hybrids (Table 2). At this hybrid zone some introgression of genic markers has been
detected, but atypical markers are found only in animals close to the zone (Bee and Close
1992). Thus despite some gene flow, each taxon is maintaining a substantial and distinct
identity (Table 1; Bee and CLose 1992) and so should be regarded as separate species.
The eastern Petrogale radiation is therefore considered to comprise eight species:
P. godrnani (containing only the godmani chromosome race), P. assirnilis (containing only
the assimilis chromosome race), P. inornata, P. herberti (containing only the herberti
chromosome race), P. penicillata (containing only the penicillata chromosome race) and three
un-named species, Petrogale 'Cape York', Petrogale 'Mareeba' and Petrogale 'Mt Claro'.
Species Accounts
The principle means of species identification used in this revision is chromosome number
and morphology. However, the genic data have been useful in establishing whether each
taxon should be regarded as a separate species. We fully realise that this account does not
provide a ready classification for naturalists or field biologists, this being a common but obvious
problem with the identification of cryptic species. However, even in the absence of chromo-
some data, many Petrogale specimens can be provisionally identified on geographical
grounds, although this means of identification will be limited to areas in which karyotypic
studies have already been conducted.
Petrogale penicillata Gray
Brush-tailed rock wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Whereabouts of holotype or type locality unknown. Original description based on a drawing (Gray
1825); see Calaby and Richardson (1988).
Diagnosis
Karyotype: diploid number 2n =22; autosomes 1, 2, 3, 4, 5, 6 , 7, 8, 9 and 10 acrocentric;
X-chromosome also acrocentric (Fig. 1). See Eldridge et al. (1990) for G-banded karyotype
and details of rearrangements.
Description
P. penicillata is a medium-large rock wallaby showing sexual dimorphism in most body
parameters (Table 4). The measurements for P. penicillata show some overlap with similar
measures for P. herberti but, on average, P. penicillata tends to be larger than P. herberti
and the other eastern taxa (Table 4). However, specimens of P. penicillata from the
Grampians, Vic., are considerably smaller than those found elsewhere in south-eastern
Australia (Close et al. 1998).Table 3. Summary of eastern Petrogale taxonomy from 1888 to present
+, indicates that the taxon was not yet known; -, indicates that the taxon was known, but not examined
Petrogale Petrogale Petrogale Petrogale Peirogale Petrogale Petrogale Petrogale Petrogale
penicillata herberti inornata assimilis puella mareeba, sharmani, godmani coenensis,
(Gray, 1825) Thomas, 1926 Gould, 1842 Ramsay, 1877 Thomas, 1926 sp. nov. sp. nov. Thomas, 1923 sp. nov.
Thomas 1888 P. penicillata P. inornata P. penicillata
Iredale and Troughton P. penicillata P. inornata P. inornata P. inornata P. inornata + P. inornata
1934 herberti inornata inornata puella godmani
Tate 1948 P. penicillata P. penicillata P. inornata P. inornata P. inornata + P. inornata
penicillata herberti inornata inornata puella godmani
Marlow 1965 P. penicillata P. penicillata P. inornata P. inornata P. inornata + P. inornata
penicillata herberti inornata inornata puella godmani
Troughton 1967 P. penicillata P. inornata P. inornata P. inornata P. inornata + P. inornata
herberti inornata assimilis puella godmani
Ride 1970 P. penicillata P. penicillata P. penicillata P. penicillata P. pencillata + P. godmani
penicillata herberti inornata inornata (inornata)
Poole 1979 P. penicillata P. penicillata P. penicillata P. penicillata P. pencillata P. penicillata P. penicillata P. penicillata
penicillata herberti inornata assimilis puella ssp. nov. 3 ssp. nov. 4 godmani
Briscoe et al. 1982 P. penicillata P. penicillata P. inornata P. inornata P. inornata P. inornata P. inornata P. godmani P. godmani
penicillata herberti (inornata (assimilis (puella (Mareeba (Mt Claro (godrnani (Cape York
race) race) race) race) race) race) race)
Strahan 1983 P. penicillata P. penicillata P. inornata P. inornata P. inornata P. inornata P. inornata P. godmani P. godmani
penicillata herberti inornata assimilis puella Mareeba Mt Claro godmani Cape York
race race race
Calaby and Richardson P. penicillata P. penicillata P. inornata P. assimilis P. assimilis P. assimilis P. assirnilis P. godmani P. godmani
1988
Sharman et al. 1990 P. penicillata P. penicillata P. inornata P. assimilis Mareeba race Mt Claro race P. godmani P. godmani
penicillata herberti Cape York
race
This paper P. penicillata P. herberti P. inornata P. assimilis P. assimilis P. mareeba P. sharrnani P. godmani P. coenensisTaxonomy of Rock Wallabies. I.
Fig. 1. Idiogram of karyotypes for the eastern Petrogale species.
Except where chromosomes are polymorphic, only one homologue from
each chromosome pair is shown.
Typically, P. penicillata is brown above, tending to rufous on the rump and grey on the
shoulders. The chest and belly are paler. In some animals a white blaze is present on the
chest. White to buff cheek stripe, black dorsal stripe from forehead to back of head.
Exterior of ears black, inside of ears buff. Black axillary patch often extending as a dark
stripe to margin of hind legs. A pale grey side-stripe is sometimes present. Feet and paws
dark brown to black. Tail darkens distally with a prominent brush. Especially in southern
populations the pelage is long and thick, particularly about the rump, flanks and base of
tail. Specimens from the north of the range tend to be lighter and have a less prominent
tail brush, making them similar to southern specimens of P. herberti.
Distribution
P. penicillata was formerly widespread and abundant in south-eastern Australia but
appears to have declined in many areas, including western New South Wales and Victoria
(Close et al. 1988; Short and Milkovits 1990). Despite this apparent range contraction,
P. penicillata is still the most widespread Petrogale in eastern Australia, being found inTable 4. Mean body dimensions of adult (MESIV or older) eastern Petrogale
(MES, molar eruption stage; see Sharman et al. 1964.) All measurements in rnillimetres, weight in kilograms. Range given in parentheses
Species Sex n Head Ear Arm Leg Hind foot Head and body Tail Weight
P. coenensis
P. godmani
P. mareeba
P. sharmani
P. assimilis
P. inornata
P. herberti
P. penicillataTaxonomy of Rock Wallabies. I.
scattered colonies throughout the Great Dividing Range from Nanango, 100 km north-west
of Brisbane, Queensland (where it forms a hybrid zone with P. herberti), southwards to East
Gippsland, Victoria (Fig. 2). There is also a small isolated population in the Grampians,
western Victoria (Fig. 2). Naturalised populations are found in Hawaii (Laze11 et al. 1984)
and New Zealand (Wodzicki and Flux 1967). The collection localities of all specimens with
the diagnostic P. penicillata karyotype are shown in Fig. 2. Other populations attributable
to P. penicillata are known (e.g. see Short and Milkovits 1990) but are yet to be examined
cytologically.
/ Adelaide
P. penicillata
Melbourne
A-
Sydney
-
Fig. 2. Distribution of P, penicillata in south-eastern Australia, showing collection
localities.
Petrogale herberti Thomas
Herbert's rocky wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Holotype. British Museum (Natural History) (BMNH) 22.12.29.9. Adult female, skin and skull,
collected by T. V. Sherrin, 2.i. 1922 (Thomas 1926).
Type locality. Eidsvold, Burnett R., south-east Queensland.
Diagnosis
Karyotype: diploid number 2n=22; autosomes 1, 2, 3, 5 , 6 , 7 , 8 , 9 and 10 acro-
centric, autosome 4 submetacentric (Fig. 1 ) ; X-chromosome usually acrocentric but can be
metacentric (Fig. 1). See Eldridge et al. (1990) for G-banded karyotype and details of
rearrangements.
Description
P. herberti is a medium-sized rocky wallaby showing sexual dimorphism in most body
parameters (Table 4 ) . The external dimensions of P. herberti show some overlap with similar
measurements from both P. penicillata and P. inornata (Table 4).M. D. B. Eldridge and R. L. Close
Typically, specimens of P. herberti are grey-brown above, darker on the face and
shoulders, tending to tawny on the rump. Chest and belly buff to white. Indistinct pale
cheek stripe, distinct black dorsal stripe from forehead to beyond shoulders. Exterior of ears
black towards the base, inside of ears buff. Blackish axillary patch. White side-stripe,
generally more distinct than that of southern P. penicillata, from axillary patch to thighs.
Fore-arms and legs brown, fore-paws and feet dark brown to black. Tail darkens distally,
brush is less prominent than that of southern P. penicillata. Specimens from the north of
range tend to be lighter and have less prominent markings and tail brush, making them
similar to southern specimens of P. inornata. Southern specimens of P. herberti are similar
to northern specimens of P. penicillata.
Distribution
This species is widespread in south-east Queensland, being found from Nanango, 100 km
north-west of Brisbane, Queensland (where it forms a hybrid zone with P. penicillata),
northwards to the south bank of the Fitzroy R., Rockhampton (where it is parapatric with
P. inornata) and west to Mt Ball (near Rubyvale) and Mt Donneybrook (near Clermont)
(Fig. 3). The collection localities of all specimens with the diagnostic P. herberti karyotype
are shown in Fig. 3.
Petrogale inornata Gould
Unadorned (or plain) rock wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Holoptype. Female, skin, collected by B. Bynoe of HMS 'Beagle' (Gould 1842). The specimen was
later reclaimed from Gould by Bynoe and is now lost (Calaby and Richardson 1988).
Type locality. Cape Upstart, Qld.
Neotype. Australian National Wildlife Collection, CSIRO Division of Wildlife and Ecology,
Canberra (ANWC) CM11637, adult male, skin and skull; field number S-694; collected by S. C. Barker
and G. B. Sharman, 4.vii.1984, Flagstaff Bay, Cape Upstart, Qld (19°46'S.,147046'E.).
Diagnosis
Karyotype: diploid number 2n =22; autosomes 1, 2, 5, 6 , 7, 8, 9 and 10 acrocentric;
autosomes 3 and 4 submetacentric (Fig. 1); X-chromosome may be acrocentric or meta-
centric (Fig. l). See Eldridge et al. (1990) for G-banded karyotype and details of
rearrangements.
Description
P. inornata is a medium-sized rock wallaby showing some sexual dimorphism in most
body parameters (Table 4). The external dimensions of P. inornata show some overlap with
similar measurements from P. herberti but, on average, P. herberti is larger (Table 4).
The measurements for P. inornata also show significant overlap with similar measurements
for the five northern species.
The general coloration of P. inornata is similar to that for P. assimilis, P. sharmani and
P. mareeba, and tends to vary according to the substrate. Most specimens of P. inornata
are grey-brown above, paler sandy brown on underparts, fore-arms and hind-legs, and
almost buff at base of tail. However, some specimens are considerably darker, tending
towards dark grey or dark brown. A pale cheek stripe and slight mid-dorsal stripe are
occasionally present. The tail darkens to almost black towards end, with a slight brush at
tip. Animals moult in autumn to predominantly grey on back and flanks, and then become
progressively more sandy through the year. The amount of ornamentation present inTaxonomy of Rock Wallabies. I.
P. inornata varies with latitude, animals in the south of range having more noticeable
markings, including a distinct head stripe and a light side stripe, making them similar to
northern specimens of P. herberti.
Distribution
This species is found in the ranges of central coastal Queensland from the north bank
of the Fitzroy R., Rockhampton (where it is parapatric with P. herberti), northwards to the
lower Burdekin-Bowen Rs, south of Townsville (where it is parapatric with P. assirnilis)
(Fig. 3). P. inornata is also found on Whitsunday I. and is parapatric/sympatric(?) with
P. persephone around Proserpine. The collection localities of all specimens with the diag-
nostic P. inornata karyotype are shown in Fig. 3.
Fig. 3. Distribution of eastern Petrogale species in Queensland, showing collection
localities.M. D. B. Eldridge and R. L. Close
Petrogale assimilis Ramsay
Allied rock wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Holotype. Female, skin and skull, collected during the Chevert expedition (Ramsay 1877). Where-
abouts of type is now unknown.
Type locality. Palm I., north of Townsville, Qld.
Diagnosis
Karyotype: diploid number 2n=20; autosomes 1 , 2, 3, 4, 5 , 7 , 8 and 9 acrocentric;
autosomes 6 and 10 fused to form a near metacentric (Fig. 1); X-chromosome may be
acrocentric or metacentric (Fig. 1). See Eldridge et al. (1988) for G-banded karyotype and
details of rearrangements.
Description
P. assirnilis is a small to medium-sized rock wallaby showing sexual dimorphism in most
body parameters (Table 4). The external dimensions of P. assirnilis show a significant
overlap with similar measurements from P. inornata although, on average, P. inornata is
larger (Table 4). The measurements for P. assimilis tend to be similar to those for the other
four north-east Queensland taxa (Table 4).
The general coloration of P. assirnilis is similar to that for P. sharrnani, P. inornata
and P. rnareeba, and tends to vary according to the substrate. Generally, specimens of
P. assirnilis are grey-brown above (but can be dark brown), paler sandy brown on under-
parts, fore-arms, hind-legs and at base of tail. Some specimens tend to russet on the rump
and base of tail. A pale cheek stripe, slight axillary patch and indistinct dorsal head stripe
are occasionally present. Paws and feet darker than limbs. Tail darkens to almost black
towards end, with a slight brush at tip. Moult as for P. inornata.
Distribution
This species is widespread in north-east Queensland, being found from Townsville south-
wards to the lower Burdekin-Bowen Rs (where it is parapatric with P. inornata), north-west
to Croydon and south-west to Hughenden and Mt Hope. P. assirnilis is also found on Palm
and Magnetic Is. The collection localities of all specimens with the diagnostic P. assimilis
karotype are shown in Fig. 3.
Petrogale sharmani, sp. nov.
Mt Claro (or Sharman's) rock wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Holotype. ANWC CM15202, adult male, skin and skull; field number S-124; collected by G . M.
Maynes and R. L. Close, 27.viii.1976.
Type locality. Mt Claro, Qld (18°52'05"S., 145°44'05"E .).
Paratype. ANWC CM15205, adult male, skin and skull; field number S-113; collected by G . M.
Maynes and R. L. Close, 21.viii.1976, Mt Claro, Qld (18°52'05"S.,145044'05"E.).Taxonomy of Rock Wallabies. I.
Diagnosis
Karyotype: diploid number 2n =20; autosomes 1 , 2, 3, 4, 6, 7, 8 and 9 acrocentric;
autosomes 5 and 10 fused to form a submetacentric (Fig. 1); X-chromosome metacentric
(Fig. 1). See Eldridge et al. (1988) for G-banded karyotype and details o f rearrangements.
P. sharmani can be distinguished from P. penicillata (2n = 22), P. herberti (2n = 22),
P. inornata (2n = 22), P. mareeba (2n = 18) and P. coenensis (2n =22) by chromosome
number and chromosome morphology (Fig. 1). P. sharmani can be distinguished from
P. godmani (2n= 20) by its possession o f a large submetacentric (5-10 fusion), whereas all
the autosomes in P. godmani are acrocentrics (Fig. 1). P. sharmani can be distinguished
from P. assimilis (2n= 20) by its possession o f a large submetacentric (5-10 fusion), whereas
P. assimilis is characterised by a smaller near metacentric (6-10 fusion) (Fig. 1).
Description
P. sharmani is a small to medium-sized rock wallaby showing some sexual dimorphism
(Table 4). The external dimensions o f P. sharmani tend to overlap with similar measure-
ments for P. assimilis, P. mareeba, P. godmani and P. coenensis (Table 4).
The general coloration o f P. sharmani is similar to that for P. assimilis, P. inornata and
P. mareeba, varying according to the substrate. Generally, specimens o f P. sharmani are
grey-brown above, paler sandy brown on underparts, fore-arms, hind-legs and at base o f
tail. Pale cheek stripe and pale patch on face between the eyes. Slight mid-dorsal head stripe
occasionally present. Tail darkening to almost black distally, with a slight brush at tip.
Moults as for P. inornata.
Distribution
Formerly known as the Mt Claro race, this species has a restricted distribution in north-
east Queensland, being found only in the Seaview and Coane Ranges, west o f Ingham
(Fig. 3). The collection localities o f all specimens with the diagnostic P. sharmani karyotype
are shown in Fig. 3.
Etymology
The specific name, sharmani, is given in recognition o f Professor G. B. Sharman and the
enormous contributions he has made to so many aspects o f marsupial biology, and for his
fundamental role in the study o f Petrogale in particular.
Petrogale mareeba, sp. nov.
Mareeba rock wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Holotype. ANWC CM1506, adult male, skin and skull; field number S-332; collected by G . M.
Maynes, 10.xi.1979.
Type locality. Mungana trucking yards, 16 km west of Chillagoe, Qld (17°06'S.,144023'E.).
Diagnosis
Karyotype: diploid number 2n= 18; autosomes 1 , 2, 3, 4, 7 and 8 acrocentric; autosomes
5 and 10 fused to form a submetacentric; autosomes 6 and 9 fused to form an acrocentric
(Fig. 1); X-chromosome metacentric (Fig. 1). See Eldridge et al. (1988) for G-banded
karyotype and details o f rearrangements.
P. mareeba can be distinguished from P. penicillata (2n =22), P. herberti (2n =22),
P. inornata (2n = 22), P. assimilis (2n = 20), P. sharmani (2n = 20), P. godmani (2n = 20)
and P. coenensis (2n = 22) by chromosome number and chromosome morphology (Fig. 1).M. D. B. Eldridge and R. L. Close
Description
P. mareeba is a small to medium-sized rock wallaby showing some sexual dimorphism
(Table 4). The external dimensions of P. mareeba tend to show a significant overlap
with similar measurements from P. assimilis, P. sharmani, P. godmani and P. coenensis
(Table 4).
The general coloration of P. mareeba is similar to that for P. sharmani, P. inornata and
P. assimilis, and varies according to the substrate. Typically, specimens of P. mareeba are
grey-brown above, paler sandy brown-buff on underparts, fore-arms, hind-legs and at the
base of the tail. Some specimens are considerably darker and may be dark brown or almost
black. A pale cheek stripe and a mid-dorsal head stripe are occasionally present. Tail
darkens distally, with a slight brush at tip. A dirty white tail tip of variable length is
occasionally found in some norther populations. Moult as for P. inornata.
Distribution
Previously known as the Mareeba race, this species is found from Mareeba northwards
to the Mitchell R. and near Mt Carbine (where it forms a hybrid zone with P. godmani),
west to Mungana and south to the Burdekin R. near Mt Garnet (Fig. 3). The collection
localities of all specimens with the diagnostic P. mareeba karyotype are shown in Fig. 3.
Etymology
This species is named after the north Queensland town of Mareeba, 25 km west of
Cairns, which at the time the Mareeba race was named (Briscoe et al. 1982) was at the
centre of the known distribution.
Petrogale godmani Thomas
Godman's rock wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Holotype. BMNH 23.1.5.19. Adult male, skin and skull, collected by T. V. Sherrin, 20.vii.1922
(Thomas 1923).
Type locality. Black Mtn, 16 miles (25.7 km) south-west of Cooktown, Qld.
Diagnosis
Karyotype: diploid number 2n=20; autosomes 1 , 2 , 3 , 4, 5 , 7, 8 and 9 acrocentric;
autosomes 6 and 10 fused to form an acrocentric (Fig. 1 ) ; X-chromosome may be acro-
centric or metacentric (Fig. 1). See Eldridge et al. (1989) for G-banded karyotype and details
of rearrangements.
Description
P. godmani is a small to medium-sized rock wallaby showing sexual dimorphism in
most body parameters (Table 4). The external dimensions of P. godmani tend to overlap
with similar measurements from P. assimilis, P. mareeba, P. sharmani and P. coenensis
(Table 4).
The general coloration of P. godmani is similar to that of P. sharmani, P. inornata,
P. assimilis and P. mareeba, and varies according to the substrate. However, specimens of
P. godmani are frequently more distinctly buff or pale cinnamon on the foreparts and the
distal to of the tail is frequently dirty white.Taxonomy of Rock Wallabies. I.
Distribution
This species is found from near Mt Carbine and the Mitchell R. (where it forms a
hybrid zone with P. mareeba) northwards to Bathurst Head and west to 'Pinnacles' (Fig. 3).
The collection localities of all specimens with the diagnostic P. godmani karyotype are
shown in Fig. 3.
Petrogale coenensis, sp. nov.
Cape York rock wallaby
For details of nomenclatural history and synonymies see Table 3.
Type Data
Nolotype. ANWC CM10493, adult male, skin and skull; field number S-441; collected by T. Bush
and L. R. McQuade, 15.ix.1981.
Type locality. 'Twin Humps', north of Coen, Qld (13"47'27"S., 143O04'24"E.).
Paratypes. ANWC CM10494, adult male, skin and skull; field number S-442; collected by T. Bush
and L. R. McQuade, 15.ix.1981, at 'Twin Humps', north of Coen, Qld (13°47'27"S.,143004'24"E.).
ANWC CM13479, adult female, skin and skull; field number S-865; collected by S. C. Barker and
R. L. Close, 16.vii.1985, Edward River Rd, 10 km west of Musgrave, Qld (14"48'W'S.,143"25'34'%.).
ANWC CM13480, adult female, skin and skull; field number S-866; collected by S. C. Barker and
R. L. Close, 16.vii.1985, Edward River Rd, 10 km west of Musgrave, Qld (14°48'WS.,143025'34"E.).
ANWC CM13481, adult male, skin and skull; field number S-868; collected by R. L. Close and A. A.
Gooley, 28.vii.1985, Fall Creek Stn, south of Pascoe R., 300 m east of Kennedy Rd, Iron Range,
Qld (12°55'3U'S.,143001'3U'E.).
Diagnosis
Karyotype: diploid number 2n =22; autosomes 1, 2, 3, 5, 6 , 7, 8, 9 and 10 acrocentric;
autosome 4 submetacentric (Fig. 1); X-chromosome metacentric (Fig. 1). See Eldridge et al.
(1989) for G-banded karyotype and details of rearrangements.
P. coenensis can be distinguished from P. assimilis (2n = 20), P. sharmani (2n = 20),
P. mareeba (2n = 18) and P. godmani (2n = 20) by chromosome number and chromosome
morphology (Fig. 1). P. coenensis can be distinguished from P. penicillata (2n =22) by
its possession of a single submetacentric (chromosome 4), whereas all the autosomes in
P. penicillata are acrocentrics (Fig. 1). P. coenensis can be distinguished from P. inornata
(2n = 22) by its possession of a single submetacentric (chromosome 4), whereas P. inornata
is characterised by two submetacentrics (chromosomes 3 and 4) (Fig. 1). P. coenensis like
P. herberti (2n = 22) is characterised by a single submetacentric chromosome (chromosome
4); however, the submetacentric 4 of P. coenensis is more metacentric than that 4 of
P. herberti (Fig. 1).
Description
The data for P. coenensis are limited as only three MES IV specimens are known
(Table 4). Body dimensions for these individuals fit within the range of similar measurements
for P. godmani, P. sharmani, P. assimilis and P. mareeba (Table 4).
P. coenensis appears similar in pelage to P. godmani. Generally, specimens are grey-
brown above, with paler (sandy brown to buff) underparts, fore-arms, hind-legs and base
of tail. Pale to buff cheek stripe. Mid-dorsal head stripe extending down neck to upper
back in some specimens. Tail darker than body towards base with a slight brush at tip.
An increasing density of dirty-white hairs towards tip of tail results in a distinct silvery
tail tip in most specimens. In CM13479 fur on underparts was very short, dense and pale
(almost white), with mottled darker patches on belly and towards the rump and flanks.
CM13481 had purple coloration over the chest and throat and the distal $ of the tail was
silvery white.M. D. B. Eldridge and R. L. Close Distribution Previously known as the Cape York race, this species is apparently restricted to a small area of eastern Cape York Peninsula, from Musgrave north to the Pascoe R. (Fig. 3). The collection localities of all specimens with the diagnostic P. coenensis karyotype are shown in Fig. 3. This species may be uncommon as only small scattered colonies have been located. No signs of Petrogale were found in an area 70 km wide separating P. godmani and P. coenensis. Etymology This species is named after the north Queensland town of Coen, which is located at the centre of this species' unknown distribution and is also near the type locality. Acknowledgments This work is based on Petrogale specimens collected as part of the continuing rock wallaby project at Macquarie University, and hence we are extremely grateful to all those who have been involved during the many phases of the project. We are especially grateful to Professor Geoff Sharman and Dr Gerry Maynes, who did much of the initial work on which this revision is based. We thank Dr John Calaby and John Wombey (CSIRO Wildlife and Ecology, Canberra) for their assistance and for allowing us access to the specimens in the Australian National Wildlife Collection. We also thank the National Parks and Wildlife Services of New South Wales and Queensland, as well as the Department of Conservation and Environment, Victoria, for granting us permission to collect specimens. We thank Jenny Norman for photography, Barbara Duckworth for assistance with the preparation of figures and Dr Peter Johnston, Dr George McKay and Jane Bell for their comments on the manuscript. We are especially grateful to Dr John Calaby for his advice and encouragement. This work was supported by research grants from the Australian Research Council, Macquarie University and the University of Western Sydney, Macarthur. This is publication No. 131 of the Research Unit for Biodiversity and Bioresources, Macquarie University. References Barker, S. C., and Close, R. L. (1990). Zoogeography and host associations of the Heterodoxus octoseriatus group from rock wallabies (Macropodidae: Petrogale). International Journal of Parasitology 90, 2180-7. Baverstock, P. R., Gelder, M., and Jahnke, A. (1983). Chromosome evolution in Australian Rattus. G-banding and hybrid meiosis. Genetica 60, 93-103. Baverstock, P. R., Adams, M., and Watts, C. H. S. (1986). Biochemical differentiation among karyotypic forms of Australian Rattus. Genetica 71, 11-22. Bee, C. A., and Close, R. L. (1992). Mitochondrial DNA analysis of introgression between adjacent taxa of rock-wallabies Petrogale species (Marsupialia :Macropodidae). GeneticalResearch, in press. Briscoe, D. A., Calaby, J. H., Close, R. L., Maynes, G. M., Murtagh, C. E., and Sharman, G. B. (1982). Isolation, introgression and genetic variation in rock-wallabies. In 'Species at Risk: Research in Australia'. (Eds R. H. Groves and W. D. L. Ride.) pp. 73-87. (Australian Academy of Science: Canberra.) Calaby, J. H., and Richardson, B. J. (1988). Macropodidae. In 'Zoological Catalogue of Australia. 5. Marsupialia'. (Ed. D. W. Walton.) pp. 60-80. (Australian Government Publishing Service: Canberra.) Close, R. L., and Lowry, P. S. (1990). Hybrids in marsupial research. Australian Journal of Zoology 37, 259-67. Close, R. L., Ingleby, S., van Oorschot, R. A. H., Gooley, A. A., Briscoe, D. A., and Sharman, G. B. (1988). Identification of rock-wallabies, Petrogalepenicillata (Gray 1825), from the Grampians, Victoria, and comparison with conspecifics by examination of chromosomes, blood proteins, cell surface antigens, parasites and morphology. Australian Journal of Zoology 36, 99-1 10.
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(Angus and Robertson: Sydney.)M. D. B. Eldridge and R. L. Close Sharman, G. B., and Maynes, G. M. (1983~).Short-eared rock-wallaby Petrogale brachyotis. In 'The Australian Museum Complete Book of Australian Mammals'. (Ed. R. Strahan.) p. 221. (Angus and Robertson: Sydney.) Sharman, G. B., Frith, H. J., and Calaby, J. H. (1964). Growth of the pouch young, tooth eruption and age determination in the red kangaroo, Megaleia rufa CSIRO Wildlife Research 9, 20-49. Sharman, G. B., Close, R. L., and Maynes, G. M. (1990). Chromosome evolution, phylogeny and speciation of rock-wallabies (Petrogale :Macropodidae) . Australian Journal of Zoology 37, 35 1-63. Short, J., and Milkovits, G. (1990). Distribution and status of the brush-tailed rock-wallaby in south- eastern Australia. Australian Wildlife Research 17, 169-79. Strahan, R. (1983). (Ed.) 'The Australian Museum Complete Book of Australian Mammals.' (Angus and Robertson: Sydney.) Tate, G. H. H. (1948). Results of the Archbold Expeditions. No. 59. Studies on the anatomy and phylogeny of the Macropodidae (Marsupialia). Bulletin of the American Museum of National History 91, 239-35 1. Thomas, 0. (1888). 'Catalogue of the Marsupialia and Monotremata in the collection of the British Museum (Natural History).' (British Museum: London.) Thomas, 0. (1923). Exhibition of a new rock kangaroo from northern Queensland. Proceedings of the Zoological Society of London 1923, 177-8. Thomas, 0. (1926). On various mammals obtained during Capt. Wilkins's Expedition in Australia. The Annals and Magazine of Natural History 17, 625-35. Troughton, E. Le G. (1967). 'Furred Animals of Australia.' (Angus and Robertson: Sydney.) Wiley, E. 0. (1981). 'Phylogenetics. The Theory and Practice of Phylogenetic Systematics.' (John Wiley & Sons: New York.) Wodzicki, K., and Flux, J. E. C. (1967). Guide to the introduced wallabies in New Zealand. Tuatara 15, 47-59.
Taxonomy of Rock Wallabies. I.
Appendix. Proteins examined in electrophoretic study
Abbreviation Protein E.C. Number
AAT Aspartate aminotransferase
ACON Aconitase
ACP Acid phosphatase
ADA Adenosine deaminase
ADH Alcohol dehydrogenase
AK Adenylate kinase
ALB Albumin
CA Carbonic anhydrase
CAT Catalase
ENOL Enolase
FUM Fumarate hydratase
a-GAL a-Galactosidase
GDA Guanine deaminase
GDH Glutamate dehydrogenase
a-GPD Glycerol-3-phosphate dehydrogenase
GPD Glucosed-phosphate dehydrogenase
GPT Alanine aminotransferase
IDH Isocitrate dehydrogenase
LDH Lactate dehydrogenase
MDH Malate dehydrogenase
ME Malic enzyme
MPI Mannose-phosphate isomerase
PEP Peptidases
6PGD 6-phosphogluconate dehydrogenase
PGI (GPI) Glucose-phosphate isomerase
PGK Phosphoglycerate kinase
PGM Phosphoglucomutase
PK Pyruvate kinase
SDH (SORDH) Sorbitol dehydrogenase (L-iditol dehydrogenase)
SOD Superoxide dismutase
TF Transferrin
XDH Xanthine dehydrogenase
Manuscript received 21 February 1992; accepted 10 September 1992You can also read
