Content uploaded by Michael Archer
Author content
All content in this area was uploaded by Michael Archer on Feb 08, 2016
Content may be subject to copyright.
New Miocene Icarops material (Microchiroptera: Mystacinidae) from
Australia, with a revised diagnosis of the genus
SUZANNE HAND, MICHAEL ARCHER & HENK GODTHELP
HAND, S.l., ARCHER, M. & GODTHELP, H., 2001:12:20. New Miocene lcarops material
(Microchiroptera: Mystacinidae) from Australia, with a revised diagnosis of the genus. Memoirs
of the Association of Australasian Palaeontologists 25,139-146. ISSN 0810-8889
New fossil material referable to Icarops paradox Hand et al., 1998 is described from the early
Miocene Judith's Horizontalis Site in the Riversleigh World Heritage Property of northwestern
Queensland. Fused dentaries contain the partial lower dentition of I. paradox. The diagnosis of
the genus Icarops is revised. The new material confirms the identity of Icarops species as
mystacinids and enables re-examination of interrelationships between extinct and extant members
of this Gondwanan bat family.
S.J: Hand, M. Archer* & H. Godthelp, School of Biological Science, University of New South
Wales, New South Wales, 2052; * also Australian Museum, 6-8 College St, Sydney, New South
Wales, 2000. Received ]4 December 2000
Keywords: Mystacinidae, Icarops, Mystacina, bat, lower dentition, Miocene, Riversleigh
QMF refers to specimens held in the fossil
collections of the Queensland Museum, Brisbane.
SYSTEMAllC PALAEONTOLOGY
OrderCIllROPTERAB1wnenbach, 1779
Suborder MICROCIllROPTERA Dobson, 1875
Superfamily NocmIoNoIDEA Van Va1en,
1979
Family MYSTACINIDAE (Gray, 1843)
Icarops Hand et al., 1998
Type species. Icarops breviceps Hand, Murray,
Megirian, Archer & Godthelp, 1998
Revised diagnosis. Species of Icarops differ from
Mystacina species in the following combination
of features: It width and length approximately equal
(i.e. not especially wide); p 4 less than half MI
length, with two roots oriented longitudinally or
only slightly obliquely with respect to the
toothrow; all cusps of lower molars including
entoconids taller and more distinct, with MI-2
talonid wider than trigonid.
Other species. Icarops aenae Hand et al., 1998;
Icarops paradox Hand et al., 1998
THE FIRST pre-Pleistocene record for the
Mystacinidae and first record of this bat family
from outside New Zealand were reported by Hand
et al. ( 1998) from Miocene sediments in Australia.
Three species of the new mystacinid genus
Icarops were described: Icarops breviceps from
the middle Miocene Bullock Creek deposit of the
Northern Territory; I. aenae from the early
Miocene Wayne's Wok deposit, D Site Plateau,
Riversleigh, northwestern Queensland; and I.
paradox from the early Miocene Neville's Garden
Site, D Site Plateau, Riversleigh. The Australian
mystacinids were described mainly on the basis
of dentaries; only I. breviceps was represented
also by teeth: an M2 and MJ preserved in the
holotype.
Additional mystacinid material has since been
collected from another early Miocene deposit on
Riversleigh 's D Site Plateau, Judith 's Horizontalis
Site. This material includes a mandible with fused
dentaries containing the partial lower dentition
for a small species of Icarops. Based on
comparisons of size and alveolar patterns, the new
material is referable to I. paradox. Icarops paradox
is one of 44 species of fossil microchiropterans
recorded from Riversleigh's Oligo-Miocene and
Pliocene limestone deposits (Archer et al. 1994,
Hand 1999).
In this paper, the new Icarops material is
described and the diagnosis of the genus revised.
Taxonomy and dental terminology follows Miller
(1907) and Hand et al. (1998). Stratigraphic
nomenclature for the Riversleigh region follows
Archer et al. (1994) and Creaser (1997). The prefix
Icarops paradox (Fig. lA-D)
Holotype. QMF20808, partial mandible preserving
fragments of left and right dentaries with alveoli
forLI"C, andP.,andRC1,P,4andM,.
Type localitY~ Neville's Garden Site, D Site
Plateau, Riversleigh World Heritage Property,
Lawn Hill National Park, northwestern
Queensland (Archer et al. 1994).
Revised diagnosis. Smaller than I. breviceps and
I. aenae, and with P 2 larger and P 4 more
transversely oriented than in I. aenae (Hand et al.
1998). Additionally, it differs from I. breviceps in
MJ being less reduced (with trigonid and talonid
of subequal width).
Description. The left and right dentaries of
QMF31561 are completely fused, with no sign of
the symphysis, and meet at an angle of
approximately 45°. The mandible's anterior
margin, in lateral profile, is receding ventrally,
rounded and without a chin process. A ventral
mandibular shelf extends posteriorly to the
alveolus for the anterior root of P 4. A small but
deep invagination in the shelf's posteroventral
face marks the attachment point for the digastric
muscle/s (in the holotype of I. paradox,
QMF20808, the site for attachment of the digastric
muscles was not clear). The deI!tary decreases in
depth only slightly from P 2 to below the posterior
root of MJ. The ascending ramus, condyle and
angular process are not preserved.
Unlike other Icarops specimens (including the
holotype of I. paradox), there is no mental foramen
beneath P 2 nor further posteriorly. There is,
however, a small, dorsally directed foramen below
and between the alveoli forC1 and 11. This foramen
appears to be homologous with that found in this
region in other Icarops specimens. The incisor
alveolar region is well preserved and contained a
single pair of incisors. The canine alveolus is large
and oval. The single alveolus for Pis smaller than
the canine alveolus. As in the holotype, there is
New material. QMF31561, a partial, fused
mandible with left dentary preserving C1 and MI-3
and alveoli for I1 and p 24' and right dentary with
broken p 4' MI-3 and associated RIl.
Locality, age and lithology. Judith's Horizontalis
Site, D Site Plateau, Riversleigh (see Creaser 1997),
a freshwater limestone deposit interpreted as early
Miocene, based on stratigraphy and contained
local fauna {Archer et al. 1994: Creaser 1997).
Associated fauna. Other vertebrates recovered so
far from the Judith 's Horizontalis Site include:
frogs, lizards, turtles, passerine birds, bandicoots,
a marsupial mole, yalkaparidontid, burramyid
possum, balbarid kangaroo, carnivorous
kangaroo, and a megadermatid bat (A. Gillespie,
pers. comm. ).
evidence of crowding of teeth with overlapping
of the alveoli. Most of the crown ofP 4 is broken
offbut it has two roots oriented slightly obliquely
with respect to the toothrow; the anterior alveolus
is significantly smaller than the posterior one and
displaced towards the buccal margin of the
toothrow. The anterior alveolus for MI is also
slightly buccally displaced.
The lower dental formula is 11' CI' P 2.4' M123.
The alveoli for the canines (but not the mciso"rs;
contra Hand et at. 1998) are ventrally displaced
with respect to the toothrow. The margins of the
canine alveoli are smooth and complete. The
posterior alveolus of P 4 is compressed by the
anteriorly inclined alveolus for the anterior root
ofMI. The posterior alveolus for MI is larger than
the anterior one.
RI1 was found within the jaw but was removed
during preparation of the mandible. It is relatively
large, and deeply and evenly trifid. The crown is
not markedly extended backwards; its width and
length are approximately equal.
Cl is simple in form; its posterior base has a
small, rounded median cusp, marked by a small
notch for the anterior edge of P 2. The basal
cingulum is almost complete except at its most
anterolingual point; its anterolingual edge has a
distinct convexity just above the level of the
incisors; here the cingulum is broken and
connected to a short crest on the lower part of the
crown.
Most of the crown ofRP 4 is missing; the roots
remain. Judging from its alveoli and remaining
crown, P 4 was longer than P 2 and less than half
the length ofMJ .Its two roots are oriented slightly
obliquely with respect to the toothrow; the
anterior alveolus is smaller than the posterior one
and is close to the buccal margin of the dentary.
MI has two roots and five distinct cusps, the
hypoconulid being a small cingular cusp.
Although individualised, the cusps appear
relatively low and inclined rather than tall and
upright. The trigonid is conspicuously narrower,
but approximately the same length as the talonid.
The protoconid and hypoconid are the dominant
cusps in height and volume, but they are not
massive. The protoconid is only just taller than
the hypoconid which is taller than the metaconid;
the entoconid and paraconid are subequal in
height; all are much taller than the hypoconulid.
The protoconid shows conspicuously more wear
than the other cusps. The paracristid is just longer
than the metacristid; the protoconid contributions
to the paracristid and metacristid are longer than
the paraconid and metaconid contributions; the
metaconid and protoconid contributions of the
metacristid are more equal and meet at a more acute
angle. The cristid obliqua, in occlusal view, is
AAP Memoir 2
Fig. l.lcaropsparadox Hand, Murray, Megirian, Archer & Godthelp, 1998. QMF31561, from the early
Miocene Judith's Horizontalis Site, Riversleigh World Heritage Property, Lawn Hill National Park, northwestern
Queensland. A, lateral view; B, oblique occlusal view; C, stereopair, occlusal view. Scale bar indicates 2 mrn. v=
ventral mandibular shelf.
uncurved and contacts the trigonid at a point between the hypoconid and entoconid. A pre-
directly below the junction of the components of entocristid, straight and steeply dipping, links the
the metacristid. In lateral view there is an inflexion entoconid to the trigonid at the base of the
along the cristid obliqua close to the trigonid. The metaconid (making an angle with the metaconid
hypocristid extends from the hypoconid directly of just less than 900). The angle between the para-
to the entoconid, almost perpendicular to the axis and metacristids is relatively broad, at
of the toothrow, isolating the small hypoconulid approximately 750. The cristid obliqua and
and thereby exhibiting the myotodont condition paracristid are almost parallel to each other. There
(Menu & Sige 1971 ). The greatly bowed inflexion is a uniform, non-sinuous, continuous anterior,
in the hypocristid occurs approximately midway buccal and posterior cingulum, terminated at its
posterolingual end well short of the hypoconulid
providing a notch for the anterior cingulum of
MrArelatively strong but short lingual cingulum
occurs at the base of the trigonid.
M2 is described insofar as it differs from MI.
M2 is narrower and shorter in length but taller
than MI. The trigonid and talonid are wider but
slightly shorter, the protoconid is taller (and not
as worn) but the hypoconid is of similar size, and
the metaconid and protoconid contributions of
the metacristid are more equal and meet at a more
acute angle. The angle made between the
paracristid and metacristid is approximately 600.
The short lingual cingulum at the base of the
trigonidisweakerinM (andM ).
M3 is described insofar as it differs from M1-r
It is a slightly narrower, shorter tooth. The trigonid
is similar in width to the talonid, and the talonid is
longer than the trigonid. The protoconid is the
tallest and most massive cusp, the paracristid is
longer than the metacristid, the protoconid
contribution to the paracristid being particularly
long. All cusps are present, including the
hypoconulid.
Measurements. Maximum dentary depth (below
alveolus for PJ 1.30 mm, CI-M3length 5.65 mm,
MI-31ength 3.83 mm.
COMPARISONS
Comparisons with other mystacinids. Icarops
paradox differs from the Australian Miocene I.
breviceps and I. aenae in being smaller, with M}
less reduced (with trigonid and talonid of subequaJ
width) than in at least I. breviceps; p 21arger and p 4
more transversely oriented than in I. aenae. As in
other Icarops species, the area of attachment for
digastric muscle(s) appears to be well developed
in this specimen of I. paradox (see Hand et al.
1998). Unlike the holotype QMF20808 (and an other
extinct and extant bats examined), QMF31561 has
no mental foramen; the condition is interpreted to
be an aberrant condition in this individual.
Icarops paradox and New Zealand's
Quaternary Mystacina species (i.e., M.
tuberculata and the recently extinct M. robusta )
share the following dental and mandibular
features: a fused dentary symphysis; a single trifid
lower incisor; CI with posterior heel and
posterobasal cusp; and myotodont lower molars.
Icarops paradox differs from Mystacina species
in the following features: II width and length
approximately equal (i.e. not especially wide); far
less massive canine with shorter heel, less
developed posterobasal cusp and better
developed buccal cingulum; p 41ess than half MI
length with its two roots oriented slightly
obliquely to the toothrow; all cusps of lower
Comparisons with noctilionoids. Icarops species
share with South America 's noctilionids a deep
and robust dentary with tall ascending ramus (as
in I. breviceps) and fused dentary symphysis, a
single pair of lower incisors, two anterior
premolars, and myotodont lower molars (Hand et
al. 1998). However, noctilonids differ strikingly
from Icarops (and Mystacina) species in having
M2 with extremely wide talonid, wall-like pre-
entocristid and cristid obliqua extending to the
lingual margin of the crown, more obliquely
oriented P 4' and small, lingually displaced P .The
new Icarops paradox material additionally Jiffers
in its C1 with posterobasal cusp, and the single
lower incisor being trifid rather than bifid.
The speciose bat family Phyllostomidae
exhibits an enormous range of dental
morphologies which reflects a great variety of
diets (including blood, nectar, fruit, flesh and
insects) making phyllostomids much more difficult
to characterise than other bat groups. However,
Icarops species appear to differ from
phyllostomids (and mormoopids ) in the following
combination of features: reduction of the lower
incisors to one pair, fusion of dentary symphysis,
molars, including entoconids, taller and more
distinct, MI-2 talonid wider than trigonid.
Comparisons with molossids. Icarops species
share with many molossids (Hand et al. 1998): a
deep and robust dentary with mental foramen
occurring beneath p 2; a single pair of lower
incisors; two anterior premolars overlapping so
that their roots are oblique in the toothrow; general
appearance of MI and M2, notably myotodonty,
talonid wider than trigonid, tall hypoconid,
paraconid development similar to metaconid (i.e.
without strong reduction; except in the molossid
Cheiromeles torquatus), cristid obliqua complete
(Sige 1985, p. 170-1). Additional dental features
now found to be similar to molossids include C(
being simple in form without a long heel, and p 4
being smaller than MI.
Dental features of Icarops paradox now known
to differ from molossids include: a trifid rather than
bifid single incisor; CI with posterobasal cusp; p 2
less reduced than in most molossids (although in
molossids it is double-rooted even when very
reduced e.g. molossines, cheiromeles etc.); no
sinuous buccal cingulum in MI-J' talonid not much
wider than trigonid, without pronounced buccal
bulging of the protoconid and hypoconid; less
hypsodont. Also unlike Icarops species, many
molossids (e.g. Cheiromeles) have nyctalodont
or submyotodont lower molars, and in
Cheiromeles and many molossines MJ is more
reduced.
t\AP Mernoi
relatively large p 2 with small p 4' and myotodont
lower molars. Icarops species share with many
phyllostomids and mormoopids trifid rather than
bifid lower incisors.
Comparisons with nataloids s. I. Natalids,
thyropterids, furipterids and myzopodids are
grouped together here in the superfamily
Nataloidea following Simmons (1998). Also
grouped here are the kerivoulines (species of
Kerivoula and Phoniscus) which are generally
included in either the Natalidae or Vespertilionidae.
All taxa lack a fused dentary symphysis, retain
two or more pairs of lower incisors, and tend to
have lower molars with talonid wider than trigonid.
The lower canine is generally exceptionally
slender with a subterete shaft. Most have
nyctalodont lower molars but kerivoulines have
myotodont molars like Icarops species.
Similarities to the new Icarops paradox material
include: the presence of trifid incisors (although
lower than the metaconid. The talonid ofM3 may
be greatly or little reduced.
Comparisons with vespertilionids. Vesper-
tilionids lack a fused dentary symphysis, retain
two or more pairs of lower incisors, and tend to
have lower molars with talonid wider than trigonid.
Most groups are dominated by taxa with
nyctalodont lower molars but some ( e.g. Myotis,
Plecotus, Scoteanax, Chalinolobus) have
myotodont molars like Icarops species.
Similarities with the new Icarops paradox material
include: trifid lower incisors; p 2 and p 4 relatively
small and with p 2 singly rooted. No vespertilionids
have reduced the lower incisors to one pair, nor
have a fused dentary or an extended heel on C,.
they retain three pairs); p 2 and p 4 being relatively
small; and p 2 singly rooted.
Comparisons with rhinolophoids. Icarops
species differ from rhinolophoids in their fused
dentary symphysis, single pair of lower incisors
and in their talonid morphology. Three pairs of
lower incisors are retained by nycteridids and two
by megadermatids and hipposideriQs; these are
commonly trifid. In rhinolophoids p 2 is small, p 3 is
retained by rhinolophids and some archaic
hipposiderids, the talonid of the lower molars
tends to be rather low, narrow and short and
displays the nyctalodont pattern.
Comparisons with emballonuroids. Three pairs
of lower incisors are retained by Emballonura
and New World emballonuroid taxa, and two pairs
by Taphozous and Saccolaimus; these are
typically trifid and the outer incisor is sometimes
separated from the canine by a small gap. The
molars are usually but not always nyctalodont,
and the paraconid is reduced and conspicuously
Comparisons with extinct bat families
( archaeonycteridids, palaeochiropterygids and
hassianycteridids). The most archaic of early and
middle Eocene bats are referred to a number of
extinct families and lack most of the derived
features that distinguish bats of modern families
(see Russell & Sige 1970, Russell et al. 1973, Smith
& Storch 1981, Habersetzer & Storch 1987,
Simmons & Geisler 1998). Icarops species differ
from these early bats as follows. In
archaeonycteridids, palaeochiropterygids and
hassianycteridids (including species of
Archaeonycteris, Ageina, Australonycteris,
Icaronycteris, Palaeochiropteryx, Cecilio-
nycteris, Matthesia, Stehlinia and Hassia-
nycteris), unlike species of Icarops, the dentary
symphysis is not fused, at least two pairs of lower
incisors are present, p 3 is retained, the trigonid
cusps of the lower molars are typically much taller
than the talonid cusps, the talonid is unreduced
and hypoconulid more buccally situated, and all
cusps are more individualised. The lower molars
of most of these archaic bats exhibit the
nyctalodont pattern ( or pre-nyctalodont pattern
with the hypoconulid more buccally situated) but
some palaeochiropterygids ( e.g. in the genera
Palaeochiropteryx and Stehlinia; see Sige 1997
on the reassignment of Stehlinia to this family)
exhibit the myotodont pattern. In Icarops species
the toothrow is relatively much shorter
(foreshortened) than in archaic bats.
DISCUSSION
Comparisons of the lower dentition of Icarops
paradox with those of other microchiropterans
confirm its identity as a member of the family
Mystacinidae. Icarops paradox and New
Zealand's Quaternary Mystacina tuberculata and
M. robusta share a unique combination of dental
and dentary features including a fused dentary
symphysis; presence of a single trifid incisor; CI
with posterior heel and posterobasal cusps; an<1
myotodontlowermolars.
The new material also enables further
examination of the relationships within this
Australasian bat family. Hand et al. (1998)
suggested, on the basis of the fossil material then
known, that Icarops species were probably
plesiomorphic with respect to Quaternary
mystacinids, i.e. M. tuberculata and M. robusta.
Study of more of the lower dentition supports
this hypothesis. Icarops paradox differs from
Mystacina species in: ~ width and length being
approximately equal; L. far less massive with
shorter heel, less developed posterobasal cusps
and better developed buccal cingulum; P 41ess than
half Mi length with its two roots oriented slightly
obliquely to the toothrow; all cusps of lower
molars, including entoconids, taller and more
distinct, M talonid wider than trigonid. All of
these dentalteatures appear to be less specialised
than in New Zealand's Mystacina species,
although some ( e.g. loss of two pairs of lower
incisors, fused dentary symphysis) are more
derived than found in other microchiropteran
families including, for example, rhinolophoids,
emballonuroids, vespertilionids, archaeo-
nycteridids, palaeochiropterygids and hassia-
nycteridids.
The relationships of mystacinids to other
families of bats are less clear. Mystacina
tuberculata, a semi-terrestrial microchiropteran
endemic to New Zealand, is the sole surviving
member of the Mystacinidae. The oldest record
of the family in New Zealand is 18,000 years bp
(Mystacina spp.) from Hermit's Cave, near
Charleston, South Island, NZ (Worthy &
Holdaway 1994). Mystacina was interpreted by
Pierson et al. ( 1986), on the basis ofimmunological
distance, to be a member of the South American
superfamily Noctilionoidea, evidently dispersing
from South America to New Zealand some 35
million years ago. Subsequently, Simmons ( 1996)
found. using a 'total evidence' approach in which
morphological, reproductive, behavioural, DNA
and other data are included in a single data set,
that mystacinids are probably basal to a large
group of bats including the cosmopolitan
Vespertilionidae, Molossidae and South American
Nataloidea, implying that ancestral mystacinids
diverged from other bats at least 55 million years
ago. DNA-DNA hybridisation studies by Kirsch
et al. (1998) confirmed the affinity of Mystacina
with noctilionoids as suggested by earlier
serological studies (contra Simmons 1996) but
suggested separation from its South American
relatives occurred between 54 and 66 million years
ago. DNA sequencing studies (Kennedy et al.
1999) also indicate a mystacinid-noctilionoid
relationship, but differ from other molecular results
in placement of Mystacina within the superfamily
closer to phyllostomids and mormoopids; they
calculate the time of divergence of Mystacina from
other bat taxa to be anywhere from 45 to 68 million
years ago.
Hershkovitz (1972) and Pierson (1986) have
argued for a Southern Hemisphere origin for the
world's extant bat radiation on the basis of
distributions of endemic bat families. Sige ( 1991 )
proposed that modern bat groups evolved from
isolated immigrant archaic groups in the Southern
Hemisphere in the early Eocene. The oldest bat in
the Southern Hemisphere is a 55-million-year-old
archaeonycteridid from Australia (Hand et al.
1994), to which the much younger and more
derived mystacinids bear little resemblance. The
oldest fossil bats in South America are a late
Oligocene Brazilian molossid and middle Miocene
Colombian taxa, including the phyllostomids
Notonycteris magdalenensis and Tonatia sp., a
glossophagine, phyllostomine and Noctilio
albiventris (Czaplewski 1997).
Isolated teeth of mystacinids (yet to be
described) have now been recovered from late
Oligocene deposits at Riversleigh (and possibly
South Australia) confirming that the family
Mystacinidae was represented in Australia at least
25 million years ago. It survived here until at least
12 million years ago, occurring in middle Miocene
deposits at Bullock Creek, Northern Territory, but
had disappeared from at least the Riversleigh area
by 3-5 million years ago as evidenced by their
absence from the early Pliocene Rackham's Roost
deposit (Archer et al. 1994). The decline and,
ultimately, extinction of this lineage in Australia
may reflect the intense drying of Australia since
the late Miocene which resulted in the replacement
ofwet forests by open woodlands and grasslands
over much of the continent. The sole surviving
member of the mystacinid lineage is now restricted
to New Zealand's endemic Gondwanan-type
forests dominated by Nothofagus, Podocarpus,
Dacrydium and kauri trees (Daniel & Williams
1984).
Hand et al. (1998) have argued that the
presence of plesiomorphic mystacinids in the
Australian Tertiary record (the only pre-
Pleistocene record for mystacinids) most
parsimoniously suggests that Australia was the
immediate source of New Zealand's Quaternary
mystacinids. New Zealand separated from the rest
of Gondwana 80-90 million years ago (Fleming
1979), long before the world's first bats had
evolved. Australia, Antarctica and South America
were connected as the last fragments ofGondwana
until about 35 million years ago (Veevers 1991),
throughout the period when most modern bat
lineages first appeared in the fossil record (Sige
1991, Simmons 1996, 1998). The southern
supercontinent possibly shared a number of bat
families, but precisely when and where the curious
mystacinids originated is yet to be discovered.
A CKN O WLEDGEMENffl
Work at Riversleigh has been supported by
the Australian Research Council, Department of
the Environment, Sport and Territories, National
Estate Programme Grants (Queensland),
Queensland National Parks and Wildlife Service,
the Australian Geographic Society, Linnean
Society of New South Wales. ICI. Oueensland
AAP Memoir :
Museum, University of New South Wales and the
Australian Museum. Thanks to Anna Gillespie for
preparation of this and other Riversleigh fossil
bat specimens. The following people kindly
provided access to comparative specimens in their
institutions: A. Tennyson of the Museum of New
Zealand Te Papa Tongarewa, T. Ennis and S.
Ingleby of the Australian Museum, D. Kitchener
of the Western Australian Museum, P. Jenkins of
the British Museum (Natural History), S. Van Dyck
of the Queensland Museum, N.B. Simmons and
W. Fuchs (American Museum of Natural History).
We thank Jenni Bramrnall and Galea McGregor,
University of New South Wales, for the SEM
photographs. G. Storch and N. J. Czaplewski
critically reviewed a draft of this manuscript.
RDEHENCtS
ARCHER, M., HAND, S.I. & GODTHELP, H., 1994.
Riversleigh. Reed Books, Sydney, 264p.
CREASER, P., 1997. Oligocene-Miocene sediments of
Riversleigh: the potential significance of topography.
Memoirs of the Queensland Museum 41, 303-314.
CZAPLEWSKI, N.J., 1997. Chiroptera. 410-431 in
Kay, R.F., Madden, R.H., Cifelli, R.L. & Flynn, 1.1.
(eds), Vertebratepaleontology in the Neotropics: the
Miocene fauna of La Venta, Colombia. Smithsonian
Institution Press, Washington.
DANIEL, M. I. & WILLIAMS, G.R., 1984. A survey
of the distribution, seasonal activity and roost sites
of New Zealand bats. New Zealand Journal of
Ecology 7,9-25.
FLEMING, C.A.1979.171e Geological History of New
Zealand and its Life. Auckland University Press,
Auckland, 141p.
HABERSETZER, I. & STORCH, G., 1987.
Klassiflkation und funktione!le Fliigelmorphologie
palaogener Fledermause (Mammalia, Chiroptera).
Courier F orschungsinstitut Senckenberg 9 I, 117-
150.
HAND, S.I., 1999. Australian fossil bat diversity and
evolution. Australian Mammalogy 21, 29-32.
HAND, S.I., MURRAY, P.F., MEGIRIAN, D.,
ARCHER, M. & GODTHELP, H., 1998.
Mystacinid bats (Microchiropte'ra) from the
Australian Tertiary. Journal ofPaleontology 72,538-
545.
HAND, S.I., NOVACEK, M.I., GODTHELP, H. &
ARCHER, M., 1994. First Eocene bat from
Australia. Journal of Vertebrate Paleontology 14,
375-81.
HERSHKOVITZ, P ., 1972. The recent mammals of
the neotropical region: a zoogeographic and
ecological review. 311-432 in Keast, A., Erk, F.C. &
Glass, B. (eds), Evolution, mammals and southern
continents. State University New York Press,
Albany.
KENNEDY, M., PATERSON,A.M., MORALES, I.C.,
PARSONS, S., WINNINGTON,A.P. & SPENCER.
H.G., 1999. The long and short of it: branch lengths
and the problem ofplacing the New Zealand Short-
tailed Bat, Mystacina. Molecular Phylogenetics and
Evolution 13,405-416.
KIRSCH, I.A. W., HUTCHEON, I.M., BYRNES, O.P.
& LLOYD, B.D., 1998. Affinities and historical
zoogeography of the New Zealand Short-tailed Bat,
Mystacina tuberculata Gray 1843, inferred from
DNA-hybridization comparisons. Journal of
Mammalian Evolution 5,33-64.
MENU, H. & SloE, B., 1971. Nyctalodontie et
myotodontie, importants caracteres de grades
evolutifs chez les chiropteres entomophages.
Comptes rendu de l'Academie des Sciences, Paris
272, 1735-38.
MILLER. O.S. 1907. The families and genera ofbats.
Bulletin of the United States National Museum 57,
1-282.
PIERSON, E.D., 1986. Molecular systematics of the
Microchiroptera: higher taxon relationships and
biogeography. Unpublished PhD dissertation,
University of California, Berkeley, 262p.
PIERSON, E.D., SARICH, V.M., LOWENSTEIN,
I.M., DANIEL, M.I. & RAINEY, W.E., 1986. A
molecular link between the bats of New Zealand
and South America. Nature 6083, 60-63.
RUSSELL, D.E., LOUIS, P. & SAVAGE, D.E., 1973.
Chiroptera and Dermoptera of the French Early
Eocene. University of California Publications in
Geological Science 95,1-55.
RUSSELL, D.E. & SloE, B., 1970. Revision des
chiropteres lutetiens de Messel (Hesse, Allemange ).
Palaeovertebrata 3,83-182.
SloE, B., 1985. Les chiropteres oligocenes du Fayum,
Egypte. Geologica et Palaeontologica 19, 161-189.
SloE, B., 1991. Rhinolophoidea et Vespertilionoidea
(Chiroptera) du Chambi (Eocene inferieur de
Tunisie ). Aspects biostratigraphique, biogeo-
graphique and paleoecologique de I'origine des
chiropteres modemes. Neues Jahrbuch fUr Geologie
und Palaiintologie, Abhandlungen 182,355-376.
SloE, B., 1997. Les remplissages karstiques polyphases
(Eocene, Oligocene, Pliocene) de Saint-Maximin
(phosphorites du Oard) et leur apport a la
connaissance des faunes europeennes, notamment
pour l'Eocene moyen (MP 13).3.- Systematique:
eutheriens entomophages. 737-750 in Aguilar, l.-
P., Legendre, S. & Micaux, I. (eds), Actes du
Congres BiochroM'97, Memoires et Travaux de
l'Ecole Pratique des Hautes Etudes, 1nstitut de
Montpellier 21.
SIMMONS, N. B. 1998. A reappraisal of interfamilial
relationships of bats. 3-26 in Kunz, T.H. & Racey,
P.A. (eds), Bat biology and conservation.
Smithsonian Institution Press, Washington.
SIMMONS, N. B. 1996. Bat diversification and
palaeobiogeography: implications of anew higher-
level phylogeny. Journal ofVertebrate Paleontology
16 (3) Abstracts, 66A.
SIMMONS, N.B. & GEISLER, J.H., 1998.
Phylogenetic relationships of Icaronycteris,
Archaeonycteris, Hassianycteris, and Palaeo-
chiropteryx to extant bat lineages, with comments
on the evolution of echolocation and foraging
strategies in Microchiroptera. Bulletin of the
American Museum of Natural History 235, 1-182.
SMITH, J.D. & STORCH, G., 1981. New Middle
Eocene bats from the "Grube Messel" near
Darmstadt, W-Germany (Mammalia: Chiroptera).
Senckenbergiana biologica 61, 153-168.
VEEVERS, 1.1., 1991. Phanerozoic Australia in the
changing configuration of Proto-Pangea through
Gondwanaland and Pangea to the present dispersed
continents. Australasian Systematics and Botany 4,
1-9.
WORTHY, T.H. & HOLDAWAY, R.N., 1994. Scraps
from an owl's table -predator activity as a significant
taphonomic process recognised from New Zealand
Quaternary deposits. Alcheringa 18,229-245.
