the description of a new species of myzopoda (myzopodidae: chiroptera) from western madagascar

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(Myzopodida

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Original investigation

The description of a new species of Myzopoda(Myzopodidae: Chiroptera) from western Madagascar

By S.M. Goodman, F. Rakotondraparany and A. Kofoky

Field Museum of Natural History, Chicago, USA and WWF, Antananarivo, Departement de Biologie Animale,Universite d’Antananarivo, Antananarivo, Madagasikara Voakajy, Antananarivo, Madagascar

Receipt of Ms. 6.2.2006Acceptance of Ms. 2.8.2006

Abstract

A new species of Myzopoda (Myzopodidae), an endemic family to Madagascar that waspreviously considered to be monospecific, is described. This new species, M. schliemanni,occurs in the dry western forests of the island and is notably different in pelage coloration,external measurements and cranial characters from M. aurita, the previously described species,from the humid eastern forests. Aspects of the biogeography of Myzopoda and its apparentclose association with the plant Ravenala madagascariensis (Family Strelitziaceae) arediscussed in light of possible speciation mechanisms that gave rise to eastern and westernspecies.r 2006 Deutsche Gesellschaft fur Saugetierkunde. Published by Elsevier GmbH. All rights reserved.

Key words: Myzopoda, Madagascar, new species, biogeography

Introduction

Recent research on the mammal fauna ofMadagascar has and continues to revealremarkable discoveries. A considerable num-ber of new small mammal and primatespecies have been described in recent years(Goodman et al. 2003), and numerousother mammals, known to taxonomists,await formal description. Much of thisrecent information is based on renewedinterest in biological inventories ofthe remaining forested zones of the islandand the associated systematic studies ofspecimens obtained during these surveys. Insome cases these animals can be recognizedas new to science when first handled in thefield by specialists, while in cases of cryptic

is article as: S.M. Goodman et al.,

e: Chiroptera) from western..., Mamm

front matter r 2006 Deutsche Gesellschaft fur S

bio.2006.08.001

speciation molecular studies have been veryinformative to resolve questions of specieslimits (e.g., Olson et al. 2004; Yoder et al.2005). The bat fauna of the island is noexception – until a decade ago these animalsremained largely under studied and ongoingsurveys and taxonomic work have revealedthat they may be something on the order of30% richer in species than noted in the mostrecent treatment of this group (Peterson et al.1995). As with other mammals, in certaincases new species of bats are immediatelyrecognizable (e.g., Goodman and Cardiff2004) and in other cases molecular tools helpin defining species limits (Goodman et al.2006).

The description of a new species of Myzopoda

. biol. (2006), doi:10.1016/j.mambio.2006.08.001

augetierkunde. Published by Elsevier GmbH. All rights reserved.

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2 S.M. Goodman et al.

The chiropteran family Myzopodidae isendemic to Madagascar and as currentlyunderstood is monospecific, represented byMyzopoda aurita (Milne-Edwards andGrandidier, 1878). After the creation ofthe Myzopodidae for this species (Thomas1904), it was proposed, based on morpholo-gical grounds, to be part of the Vespertilio-noidea, which included the familiesNatalidae, Furipteridae, Thyropteridae,Vespertilionidae, Mystacinidae and Molossi-dae (Koopman 1994). More recently,using molecular studies, the Myzopodidaehas been placed in the Noctilionoidea (Teel-ing et al. 2005), which also comprises theFuripteridae, Thyropteridae, Mystacinidae,Noctilionidae, Mormoopidae and Phyllosto-midae. Only two families of bats in the worldare currently considered monospecific (Sim-mons 2005): the Craseonycteridae and theMyzopodidae.With regard to Myzopoda, Thomas (1904)referred to it as ‘‘exceedingly rare andremarkable’’, the latter point certainly asso-ciated with the large flat adhesive organs, orsuckers, attached to the thumb and hind foot;hence its vernacular name sucker-footed bat.These structures have been the subject ofdetailed morphological studies (Schliemann1970, 1971).Until about 15 years ago few observations orspecimens were available of Myzopoda. How-ever, on the basis of recent field surveys it hasbeen recorded at a variety of sites in theeastern humid forests of Madagascar (Schlie-mann and Goodman 2003; Fig. 1). There wasone previously collected specimen from Ma-hajanga, on the dry western side of the island(Schliemann and Maas 1978), whichindicated that the ecological breadth of thisgenus was greater than previously recog-nized. Over the past few years we havecaptured individuals of Myzopoda at threedifferent localities in the west. These animalsare notably different in pelage colorationthan those of the east, which lead us toinvestigate their taxonomic status. Usingdata on external, cranial and dental mor-phology we are able to document that thewestern population of Myzopoda is differentthan that of the east, and it is describedherein as a new species.

Please cite this article as: S.M. Goodman et al.,

(Myzopodidae: Chiroptera) from western..., Mamm

Material and methods

In order to examine patterns of morphologicalvariation in Myzopoda we have consulted specimenshoused in several natural history museums. Theacronyms of these institutions are: AMNH –American Museum of Natural History, New York;BMNH – The Natural History Museum, London(formerly The British Museum [Natural History]);FMNH – Field Museum of Natural History,Chicago; MNHN – Museum national d’Histoirenaturelle, Paris; PBZT – Parc Botanique et Zoolo-gique de Tsimbazaza, Antananarivo; RMNH –Rijksmuseum van Natuurlijke Histoire, Leiden;UADBA – Universite d’Antananarivo, Departe-ment de Biologie Animale, Antananarivo; UADP –Universite d’Antananarivo, Departement de Paleon-tologie, Antananarivo; USNM – National Museumof Natural History (formerly United States NationalMuseum of Natural History), Washington, DC,ZMB – Museum fur Naturkunde, HumboldtUniversitat zu Berlin [formerly Zoologisches Mu-seum], Berlin; and ZMH – Zoologisches MuseumHamburg, Hamburg. A list of Myzopoda specimensused in this study is presented as Table 1.We recorded five external measurements in milli-meters using a ruler from collected specimens beforepreparation. These included: total length, tail length,hind foot length (not including claw), ear length andforearm length. Further, we measured body mass ingrams using a spring balance. For certain specimensthese data were obtained directly from museumlabels or field catalogues, and in other cases theywere measured from liquid preserved specimens.SMG also took 10 wing and two foot measurementsusing a dial callipers from liquid preserved speci-mens: 3rd digit – metacarpal, 3rd digit – 1st phalanx,3rd digit – 2nd phalanx, 3rd digit – 3rd phalanx, 4thdigit – metacarpal, 4th digit – 1st phalanx, 4th digit –2nd phalanx, 5th digit – metacarpal, 5th digit – 1stphalanx, 5th digit – 2nd phalanx, tibia and calcar.Seven cranial or mandibular and four dentalmeasurements were made by SMG using digitalcalipers, accurate to the nearest 0.1mm: occipito-nasal length, from occipital condyles to anterior-most point of nasal bone; greatest zygomaticbreadth, width taken across zygomatic arches atthe widest point; interorbital breadth, dorsal widthat most constricted part of skull; mastoid width,greatest width across skull at mastoid processes;greatest braincase width, breadth at widest portionof braincase; lacrimal width, greatest width acrossrostrum at lacrimal projections; anterior palatalwidth (C1–C1), taken across the outer alveolarborders of the upper canines; maxillary toothrow(C1–M3), length from anterior alveolar border ofupper canine to posterior alveolar border of the


. biol. (2006), doi:10.1016/j.mambio.2006.08.001


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Fig. 1. Map of Madagascar, showing the sites specimens of Myzopoda spp. are known from and used in this study.The grey shading represents portions of the island over 1000 m in elevation. The general limits of the Central andNorthern Highlands are noted. (Map prepared by Lucienne Wilme.)

A new species of Myzopoda 3

third upper molar; upper molariform toothrow,length from anterior alveolar border of PM3 toposterior alveolar border of M3; width M3, greatestlateral-medial width of tooth; and mandible length,from the posterior-most portion of the condyles toanterior-most point of lower incisors.



Results

Using information from morphological ana-lyses we can demonstrate that the popula-tions of Myzopoda inhabiting the western


. biol. (2006), doi:10.1016/j.mambio.2006.08.001


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Table 1. List of Myzopoda specimens examined (see Material and methods for a listing of museum acronyms).

Specimens of Myzopoda auritaProvince de Fianarantsoa: Andrambovato (USNM 449282); Ifanadiana (USNM 449315); Kianjavato (AMNH

257130; FMNH 185227; UADP 226; USNM 448886, 448929, 448930, 448931, 448932, 449283, 449285);Mananjary (MNHN 1907.618)

Province de Toamasina: Ambodivoangy (MNHN 1985.434), Andasibe (FMNH 184490); Foulpointe (FMNH183990); Mahambo (RMNH 26117); Mananara (ZMB 5866); Maroantsetra (MNHN 1985.654); Tampolo (FMNH165454, 179203, 179204, 179273), ‘‘Tamatave’’ ½¼ Toamasina� (BMNH 99.11.3.5; ZMH T1488, T1489, T1840,ZMB 23584)

Province de Toliara: Analalava (USNM 577065); Bemangidy (FMNH 85237, 92832); Sainte Luce (UADBA RBJ-200); USNM 578740, 578742, 578743, 578856, 578857, 578858)

No precise locality: ‘‘Madagascar’’ (MNHN 1997.770, holotype; ZMB 22843; ZMH T1487).

Specimens of Myzopoda schliemanniProvince de Mahajanga: SF d’Ampijoroa (FMNH 177327, holotype); Ankaboka (FMNH 185790, 185791,

185793, 185794); Parc National de Namoroka (UADBA RBJ 203, RBJ 204); ‘‘Majunga ½¼Mahajanga�, sur lacote ouest’’ (MNHN 1947.292).


and eastern portions of Madagascar aredifferent from one another. Even thoughthe type specimen of M. aurita (MNHN1997.770) bears the vague designation ofbeing from ‘‘Madagascar’’, with no furtherdetails in the type description (Milne-Ed-wards and Grandidier 1878), this specimen,based on color and external measurements(see Comparisons section), is from the east.Hence, the western individuals represent anew species that is described here.

Fig. 2. Different views of holotype of Myzopodaschliemanni (FMNH 177327). Upper left – dorsal viewof cranium, upper right – ventral view of cranium andlower – lateral view of cranium and mandible.(Photograph taken by John Weinstein, Field Museumimage number Z94433-05d.)

Myzopoda schliemanni new species

Holotype: FMNH 177327, male specimenprepared as skin, skull and partial post-cranial skeleton. Field number SMG 13624.Collected by S. M. Goodman on 18 April2003 at Province de Mahajanga [Parc Na-tional d’Ankarafantsika], SF [Station Forest-iere] d’Ampijoroa, Jardin Botanique A,16119.40S, 46148.40E, 160m (Fig. 1). The skinspecimen is in good condition; although as ithad considerable subcutaneous fat, there wassome seepage of grease onto the dorsal andlower ventral pelage. The basisphenoid-ba-sioccipital suture is fully ossified and it has afull permanent dentition. The left portion ofthe rostrum is partially broken at the level ofthe lacrimal canal and the winged-portions ofthe pterygoids are broken (Fig. 2).Paratypes: Three males and four females andall are adult, except as noted. MNHN 1947.



292, female, ‘‘Majunga, sur la cote ouest’’, 12June 1940; UADBA RBJ 203 and 204, femaleand male (respectively), Province de Maha-janga, Parc National de Namoroka, An-driabe, about 5 km south of Namoroka(village), 16124030.600 and 45118039.500, 110m,


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13 October 2004; FMNH 185790, 185791,male and sub-adult female (respectively),Province de Mahajanga, Ankaboka,15102.240S, 47147.130E, 200m, 4 June 2000;FMNH 185793, 185794, male and female(respectively), Province de Mahajanga,Ankaboka, 15102.240S, 47147.130E, 200m,5 June 2000.Etymology: The species name has beenchosen to honor Professor Dr. HaraldSchliemann, University of Hamburg, inrecognition of his long interest in sucker-footed bats, including several importantanatomical studies (e.g., Schliemann 1970,1971). Professor Schliemann has recentlyretired from his university post after a longand fruitful career, including being theDirector of the Zoologisches Institut undZoologisches Museum, Hamburg. We sug-gest Schliemann’s sucker-footed bat as thevernacular name of this species and easternsucker-footed bat for M. aurita.Distribution: Myzopoda schliemanni isknown from three precise localities in thedry deciduous forest zone of western Mada-gascar from Ankaboka at the northern limitof its distribution to Namoroka at thesouthern limit (Fig. 1). These localities areapproximately 300 km apart and in lowlandhabitat. The holotype was obtained atAmpijoroa, which is also a lowland site andabout 80 km inland direct distance from thecoastal town of Mahajanga. The fourthcollection locality, of the Paris specimen(MNHN 1947. 292), is vague. On the basisof extrapolation of its preferred habitat type(see below), it is likely that this species has abroader distribution in the dry deciduouszones of the west, particularly further northto perhaps at least the Sambirano region inthe vicinity of Ambanja.Measurements of holotype: Measurementsare in mm and the body mass in g. Totallength, 107; tail length, 44; hind foot length,6; ear length, 30; forearm, 47; weight, 9.9;occipitonasal length, 14.9; greatest zygomaticbreadth, 11.7; interorbital breadth, 4.1; mas-toid width, 9.8; greatest braincase width, 8.8;lacrimal width, 5.9; anterior palatal width(C1–C1), 4.0; maxillary toothrow (C1–M3),5.9; upper molariform toothrow, 4.3; widthM3, 2.0; mandible length, 11.0; 3rd digit –



metacarpal length, 37.2; 3rd digit – 1stphalanx length, 16.5; 3rd digit – 2nd phalanxlength, 16.6; 3rd digit – 3rd phalanx length,11.7; 4th digit – metacarpal length, 36.0; 4thdigit – 1st phalanx length, 13.6; 4th digit –2nd phalanx length, 14.0; 5th digit – meta-carpal length, 37.2; 5th digit – 1st phalanxlength, 11.5; 5th digit – 2nd phalanx length,8.5; tibia length, 15.0; calcar length, 10.9.Description: Myzopoda schliemanni is arelatively small microchiropteran bat with aforearm length of 45–49mm. Like the othermember of this genus it has very prominentfunnel-shaped ears, which are not joined attheir base, and the external ear conchattaches close to the base of the mouth.There is an odd mushroom-shaped process atthe base of each ear that is probably thetragus (Thomas 1904; cf. Koopman 1994).Distinct large flat adhesive organs, or suck-ers, are attached to the thumb and hind foot.Second digit of wing reduced to the meta-carpal and a small phalanx. Tail elongatedand reaches terminus of extensive uropata-gium.The hairs of the dorsum have buff-browndistal tips merging to moderate mouse-graybases. The venter is a uniform light mouse-gray. The posterior portion of the braincasetends to be less bulbous than in M. aurita, thepalatine foramen is constricted and with anarrow ‘‘v-shape’’, the nasal sulcus is shallowand relatively broad, the nasals are narrowand rectangular in shape and the frontals arenot notably inflated. Dental formula in adultsis 2/3–1/1–3/3–3/3. It is notably smallerin several external measurements thanM. aurita.Variation: t-Tests were conducted of theexternal, cranial and dental measurementsof adult M. aurita (maximum n ¼ 36) andM. schliemanni (maximum n ¼ 7), andneither species was found to be sexuallydimorphic. In all subsequent comparisonsthe sexes are combined for each species.Considerable variation, as reflected by stan-dard deviation, was found in certain externalmeasurements for M. aurita (Tab. 2). Whenthe measurements are restricted to a singlecollector there is a notable reduction in thestandard deviation of total length and hindfoot length, while other measurements, such


. biol. (2006), doi:10.1016/j.mambio.2006.08.001


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Please cite this article as: S.M. Goodman et al., The description of a new species of Myzopoda

(Myzopodidae: Chiroptera) from western..., Mamm. biol. (2006), doi:10.1016/j.mambio.2006.08.001

Table

2.

Exte

rnal

mea

sure

men

ts(i

nm

m)

and

body

mas

s(i

ng)

ofM

yzop

oda

spp.

Sam

ples

are

ofad

ult

san

ddiv

ided

bet

wee

nea

ster

nan

dw

este

rnlo

caliti

es.

Des

crip

tive

stat

isti

cspre

sente

das

mea

n7

stan

dard

devi

atio

n,

min

imum

–m

axim

um

,n.

Sam

ples

mea

sure

dby

asi

ngle

fiel

dco

llec

tor

(SM

G)

are

init

alic

s.t-

Test

com

par

isons

are

pre

sente

dbet

wee

nth

eco

mple

tew

este

rnan

dea

ster

nsa

mple

s.M

easu

rem

ents

ofM

.au

rita

holo

type

wer

em

easu

red

direc

tly

from

liqu

idpre

serv

edsp

ecim

en.

Tota

lle

ngth

Tail

length

Hin

dfo

ot

length

Ear

length

Fore

arm

length

Body

mas

s

Myz

opod

asc

hliem

anni

Holo

type

FMNH

17732

7107

44

630

47

9.9

Com

bined

102.37

8.9

645.67

1.3

65.77

0.5

830.97

0.7

547.47

1.3

09.37

1.3

4sa

mple

s92–10

7,

n¼

344–47,

n¼

75–6,

n¼

630–32

,n¼

745–49,

n¼

77.8

–10

.3,

n¼

3

Myz

opod

aau

rita

Holo

type

MNH

N1997.

770

—51

635

48

—

Com

bined

110.37

4.4

746.97

3.0

37.47

1.7

033.97

1.4

548.87

1.3

78.67

0.8

5sa

mple

s101–1

18,

n¼

26

40–53,

n¼

36

5–101

,n¼

30

31–39

,n¼

27

46–52,

n¼

31

7.2

–10

.5,

n¼

26

112.77

1.5

347.37

3.0

65.37

0.5

832.77

1.1

547.77

1.5

29.37

0.2

5111–1

14,

n¼

344–50,

n¼

35–6,

n¼

332–34

,n¼

346–49,

n¼

39.0

–9.

5,

n¼

3

t¼

2:05,df¼

27

n.s

.t¼

2:07,df¼

32

t¼

2:03,df¼

36

n.s

.P¼

0:01

P¼

5:7�6

P¼

0:007

3rd

dig

it–

met

acar

pal

3rd

dig

it–

1st

phal

anx

3rd

dig

it–

2nd

phal

anx

3rd

dig

it–

3rd

phal

anx

4th

dig

it–

met

acar

pal

4th

dig

it–

1st

phal

anx

Myz

opod

asc

hliem

anni

Holo

type

FMNH

17732

737.2

16.5

16.6

11.7

36.0

13.6

Com

bined

36.07

1.0

816.87

0.5

718.27

0.7

411.87

1.9

135.67

1.4

414.17

0.3

9sa

mple

s34.2

–37.2

,n¼

816.1

–17.8

,n¼

816.6

–18.7

,n¼

810.6

–13.8

,n¼

733.5

–37.7

,n¼

813.6

–14.7

,n¼

8

Myz

opod

aau

rita

Holo

type

MNH

N1997.

770

38.2

18.0

20.5

12.3

38.7

14.7



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Com

bin

ed37.57

1.0

117.37

0.5

918.67

0.9

212.47

0.8

136.97

1.1

114.57

0.3

9sa

mple

s35.6

–39.0

,n¼

17

16.3

–18.6

,n¼

19

17.0

–20.5

,n¼

19

11.0

–13.7

,n¼

18

34.97

38.7

,n¼

17

13.5

–16.5

,n¼

19

t¼

2:07,df¼

23

n.s

.n.s

.n.s

.t¼

2:07,df¼

23

t¼

2:06,df¼

25

P¼

0:005

P¼

0:02

P¼

0:04

4th

dig

it–

2nd

phal

anx

5th

dig

it–

met

acar

pal

5th

dig

it–

1st

phal

anx

5th

dig

it–

2nd

phal

anx

Tibia

Calc

ar

Myz

opod

asc

hliem

anni

Holo

type

FMNH

17732

714.0

37.2

11.5

8.5

15.0

10.9

Com

bin

ed15.57

0.7

736.77

1.5

611.67

0.4

810.37

0.9

916.27

1.2

113.27

1.7

7sa

mple

s14.0

–16.6

,n¼

834.7

–38.9

,n¼

810.8

–12.4

,n¼

88.5

–11

.6,

n¼

814.8

–18.4

,n¼

810.9

–16.1

,n¼

8

Myz

opod

aau

rita

Holo

type

MNH

N1997.7

70

16.3

38.4

12.7

10.8

16.4

14.0

Com

bin

ed15.07

0.8

738.17

1.1

311.67

0.5

010.37

0.6

217.27

0.6

114.07

1.0

8sa

mple

s13.5

–16.5

,n¼

19

36.0

–40.0

,n¼

17

10.7

–12.7

,n¼

19

8.9

–11

.7,

n¼

19

16.0

–17.9

,n¼

15

12.6

–16.1

,n¼

14

n.s

.t¼

2:07,df¼

23

n.s

.n.s

.t¼

2:08,df¼

21

n.s

P¼

0:02

P¼

0:01

1Onthebasisofthis

rangeandassociatedvariance

itcanbeassumed

thatthesemeasurements

are

amixture

ofthose

withandwithoutthehindclaw.

Statisticaltestsnotconducted.



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as tail length, continue to show the same levelof variation (Tab. 2). Within M. schliemannithere is little variation in the dorsal buff-brown and ventral mouse-gray pelage colora-tion or in cranial and dental measurementsamongst the specimens obtained at severaldifferent localities. The pelage coloration ofM. aurita is a consistent dark brown.Comparisons: In the holotype skin specimenof M. schliemanni (FMNH 177327) fromAmpijoroa, the dorsum fur, which is longerand shaggier than in M. aurita, has buff-brown distal tips merging to moderatemouse-gray bases. The ventrum is a uniformlight mouse-gray (Fig. 3). In contrast, thepelage of M. aurita is characterized bybeing a uniform dark brown, bordering onblackish, on both the ventrum and dorsum(Fig. 3); although a few individuals have adark golden-brown tinge to the pelage. In thespecimens of M. schliemanni from Ankabokaand Namoroka, all of which were preservedin formaldehyde and subsequently trans-ferred to ethanol, the pelage coloration isconsistent with the holotype of M. schlie-manni and distinctly lighter than in indivi-duals of eastern M. aurita.Even though the holotype of M. aurita hasbeen stored in alcohol for over 130 years, thepelage is not completely foxed and the colorremains notably darker than individualsof this genus from western Madagascar.Further, the specimen’s external measure-ments fall within the range of the easternspecies (Tab. 2). Hence, we conclude that theholotype represents M. aurita as definedherein.The external measurements of M. schliemanniare distinctly smaller than M. aurita (Tab. 2).For total length, hind foot length, ear lengthand forearm length these differences arestatistically significant. Perhaps most notablein this regard is ear length, which shows lessoverlap between the two species than theother external measurements. Several of thewing bone measurements show statisticallysignificant differences between the two spe-cies, and in each case it is M. schliemanni thatis smaller than M. aurita (Tab. 2).The two species show broad overlap incranial and dental measurements (Tab. 3)and none of these comparisons are statisti-



cally significant. The tooth formula is iden-tical in both species. Further, no notabledifference was found in tooth or cusp shape.In M. schliemanni the cranium tends tobe slightly less bulbous in appearance thanin M. aurita. The palatine foramen inM. schliemanni is notably more constrictedand a narrow ‘‘v-shape’’ as compared to themore open and broad ‘‘U-shape’’ in M. aurita(Fig. 4). Further, there are distinct differencesin aspects of the rostral morphology in thetwo species (Fig. 5). In M. aurita the nasalsulcus is distinctly narrower and deeper thanin M. schliemanni. Further, in M. schliemannithe nasals form a largely parallel structure,while in M. aurita they are expanded poster-iorly and then constricted. The frontals arenotably more inflated in M. aurita than inM. schliemanni. The holotypes of M. schlie-manni and M. aurita were directly comparedand these differences are consistent betweenall adult specimens of the two taxa.Principal component analyses were con-ducted on external, wing bone, cranial, anddental measurements of the specimens ofMyzopoda spp. Of these four comparisons itwas only the external measurements thatshowed a clear separation of M. schliemanniand M. aurita (Fig. 6), in which the first twofactors explained 69.7% of the variance. Earlength and forearm length had the heaviestloadings (Tab. 4). These results are concor-dant with the univariate analyses.Natural history: It has been noted in theliterature that Myzopoda aurita is a denizenof moist tropical forests and is often foundassociated with the large broad-leaf plants,most notably Ravenala madagascariensis ofthe Family Strelitziaceae (Schliemann andMaas 1978; Koopman 1994; Gopfert andWasserthal 1995). This tree with banana-likeleaves, reaching 10–15m tall, often growsnaturally in areas with moist ground. Rave-nala is also one of the more importantpioneering plants in heavily degraded habi-tats, particularly the eastern biome, afterforest clearing and the successive passage offire (Blanc et al. 2003). It is presumed that thesucker-like structures on this bat’s wrists andankles are used to adhere to slick flat leaf-surfaces, aiding the bat in climbing up theleaves or adhering to a roosting position. In


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Fig. 3. Image above, photograph ventral view of holotype of Myzopoda schliemanni (FMNH 177327) obtained on18 April 2003 in the Station Forestiere d’Ampijoroa, Parc National d’Ankarafantsika. (Photograph taken by StevenM. Goodman.) Image below, photograph ventral view of typical eastern M. aurita captured at Tampolo.(Photograph taken by Achille P. Raselimanana.)



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Table

3.

Cran

ial

and

denta

lm

easu

rem

ents

(in

mm

)of

Myz

opod

asp

p.

Sam

ple

sar

eof

adult

san

ddiv

ided

bet

wee

nea

ster

nan

dw

este

rnlo

caliti

es.

Des

crip

tive

stat

isti

cspre

sente

das

mea

n7

stan

dard

devi

atio

n,m

inim

um

–m

axim

um

,n.No

signifi

cant

dif

fere

nce

sw

ere

found

bet

wee

nth

etw

oge

ogr

aphic

sam

ple

sbas

edon

t-Te

stco

mpar

isons.

Occ

ipit

o-n

asal

length

Gre

ates

tzy

gom

atic

bre

adth

Inte

rorb

ital

bre

adth

Mas

toid

wid

thGre

ates

tBra

inca

sebre

adth

Lacr

imal

wid

th

Myz

opod

asc

hliem

anni

Holo

type

FMNH

17732

714.9

11.7

4.1

9.8

8.8

5.9

Com

bined

15.17

0.2

311.57

0.1

34.17

0.0

89.37

0.5

69.07

0.1

75.97

0.1

0sa

mple

s14.9

–15.5

,n¼

611.4

–11.7

,n¼

54.0

–4.

2,

n¼

78.5

–9.

8,

n¼

78.8

–9.2

,n¼

45.7

–5.

9,

n¼

4

Myz

opod

aau

rita

Holo

type

MNH

N1997.

770

15.2

11.8

4.0

9.6

9.0

5.9

Com

bined

15.37

0.4

211.77

0.3

64.27

0.1

79.57

0.5

68.97

0.2

15.97

0.1

9sa

mple

s14.6

–16.2

,n¼

22

11.2

–12.3

,n¼

16

3.9

–4.

5,

n¼

23

8.6

–10

.3,

n¼

21

8.6

–9.2

,n¼

14

5.4

–6.

1,

n¼

16

C1–C1

C1–M

3U

ppe

rm

ola

rifo

rmW

idth

M3

Man

dib

lele

ngth

Myz

opod

asc

hliem

anni

Holo

type

FMNH

17732

74.0

5.9

4.3

2.0

11.0

Com

bined

3.87

0.1

45.97

0.1

94.27

0.1

21.87

0.0

811.37

0.2

8sa

mple

s3.6

–4.

0,

n¼

75.6

–6.

1,

n¼

74.0

–4.

3,

n¼

41.7

–2.

0,

n¼

411.0

–11.8

,n¼

7

Myz

opod

aau

rita

Holo

type

MNH

N1997.

770

3.9

5.9

4.1

1.8

11.6

Com

bined

3.97

0.1

66.07

0.1

44.27

0.1

21.87

0.0

811.67

0.3

6sa

mple

s3.6

–4.

1,

n¼

24

5.7

–6.

2,

n¼

24

4.0

–4.

5,

n¼

15

1.7

–2.

0,

n¼

16

11.0

–12.6

,n¼

19



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Fig. 4. Views of the palatine foramen in (left) Myzopoda schliemanni (Holotype FMNH 177327) and (right)M. aurita (FMNH 179273). The difference in the foramen shape in these two specimens is consistent betweenthe two species. The left pterygoid (as viewed here) of M. schliemanni is broken and bent laterally and the rightone is completely missing. (Drawing by Rebecca Kramer.)


1948, the late Harry Hoogstraal found thisspecies in a Ravenala leaf in southeasternMadagascar, in this case a young uncoiledone (Schliemann and Goodman 2003; fieldcatalogue in the FMNH), and althoughnumerous field researchers have subsequentlysearched the leaves of this plant for Myzopo-da, this is the only direct observation ofwhich we are aware to support this associa-tion. Captive individuals readily climb upRavenala leaves (Gopfert and Wasserthal1995).An interesting plant–bat relationship occursin the genus Thyroptera, a member of theNewWorld family Thyropteridae, that showsremarkable parallel development of suctiondiscs on the wrists and ankles as found inMyzopoda. Thyroptera, composed of threedistinct species, are known to use theirsuckers to grip the slick leaves of Heliconia(Family Musaceae) and Calathea (FamilyMarantaceae), which they use as day roostsin often heavily degraded forest (Findley andWilson 1974; Vonhof and Fenton 2004).Analyses of the functional morphology ofthese appendage structures and molecularstudies of the phylogenetic relationships ofThyroptera and Myzopoda (Schliemann 1970,1971; Teeling et al. 2005) reveal that theseremarkable adaptations are convergent be-tween the two genera. Even though there isonly one direct published observation of



Myzopoda roosting in Ravenala leaves, con-text and inference would imply that theaxiom about the association of this bat andplant is correct. Thus, we assume that thepreferred natural day roost sites of Myzopodaare plants with broad and smooth leaves,such as Ravenala or other plants such asPandanus (Family Pandanaceae), certainspecies of palms and ferns, introduced Musa(Family Musaceae), and Typhonodorum(Family Araceae).For the specimens of M. aurita obtained inthe eastern biome and for which we havedetails on the specific situation where theywhere captured, the majority were within orclose to stands of Ravenala and generally atelevations below 50m. In a vast portion ofthe east, from at least Vangaindrano toMaroantsetra, much of the former extensivelowland forests have been cleared and densestands of Ravenala are the dominant pioneer-ing species in the resulting secondary habitat.Myzopoda has been recorded in this zone atseveral different sites and based on extra-polation presumably has a broad distributionin heavily degraded lowland eastern areas,particularly those dominated by Ravenala.Our records of Myzopoda from westernMadagascar are associated with locally mesicconditions and for most cases in closeproximity to Ravenala or other plants withbroad and smooth leaves. The holotype of


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Fig. 5. Views of the upper rostrum in (left) Myzopodaschliemanni (Holotype FMNH 177327) and (right)M. aurita (FMNH 179273). The differences in the nasalsulcus, nasal shape and frontals in these two speci-mens are consistent between the two species.(Drawing by Rebecca Kramer.)


M. schliemanni was captured in a mist netat the edge of a marsh, near the StationForestiere d’Ampijoroa, which included Ra-venala and Raphia palms, albeit the former



not a common member of the local plantcommunity. The Ankaboka sites where thisspecies was netted contain highly degradedlowland areas and the animals were capturedover streams in gallery forest. AlthoughRavenala was not specifically noted in thiszone, it can be found in the general vicinity.Finally, at Namoroka both specimens werecaptured by hand while roosting in a cave(Kofoky et al. 2006) and an additionalindividual was netted over a stream in amixed marsh woodland area. The habitatimmediately surrounding the cave site wasdry deciduous forest, and about 750–1000mdeeper in the forest was a permanent riversystem with Raphia palms and large strap-leaf Pandanus. A third individual captured atNamoroka and released was obtained inhabitat heavily modified by human activitiesand included an extensive marsh system withRaphia, Pandanus and a few examples ofRavenala, and mango trees on slightly higherground.The holotype of M. schliemanni was obtainedon 18 April and had notably large deposits ofyellow sub-cutaneous fat. Further, the testi-cles were scrotal, measured 5� 3mm, and theepididymides were convoluted. The scrotumcoloration was noted as being, ‘‘dull yellowwith dark spots’’. Amongst the four speci-mens obtained at Ankaboka in early Junethere was one sub-adult with an unossifiedskull and partial non-permanent dentition.One of the animals handled at Namoroka inmid-October was a female with developedmammae. Nothing definitive can be statedabout the reproductive season in this species,but it would appear that it is not confined toa narrow period of the year.Several male specimens of M. aurita werenoted as having scrotal testes, including oneindividual on 14 March in which theymeasured 4� 3mm and individuals on 14April and 24 May that measured 5� 4mmand with convoluted epididymides. Thus, itwould appear that reproductive males retaintheir testicles in a descended position for atleast 3 months per year.In two cases of male M. aurita with enlargedtestes it was recorded in the field cataloguethat the scrotum was ‘‘light beige in colora-tion with black spots around testicles’’. It has


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-1.5-1.0-0.50.00.51.01.52.02.53.0

-2.5 -1.5 -0.5 0.5 1.5 2.5-2.5-2.0-1.5-1.0-0.50.00.51.01.52.02.5

-2.5

-1.5

-0.5

0.5

1.5

2.5

-2.5 -2.0 -1.5 -1.0 -0.5 0.0 0.5 1.0 1.5 2.0-2.5-2.0-1.5-1.0-0.50.00.51.01.52.0

-3.5 -2.5 -1.5 -0.5 0.5 1.5 2.5 3.5

-2.5 -1.5 -0.5 0.5 1.5 2.5

Fig. 6. Projection of factor 1 (x-axis) and factor 2 (y-axis) in principal component analysis of external (upperleft), wing bone (upper right), cranial (lower left) and dental (lower right) measurements of Myzopoda aurita(black squares) and M. schliemanni (light squares). Loading contributions of variables to each axis are shown inTable 4.


already been noted in the literature that maleMyzopodidae have pigmented portions oftheir external sexual organs (Kermott andTimm 1988), which in this case was scattereddark brown pigment to the tunica vaginalissurrounding the epididymis. The function ofthe colored testicular pouch is uncertain, butmay be associated with thermoregulation,visual communication, or a shield againstultraviolet radiation. Given that the pigmen-tation in Myzopoda forms an irregularpattern, rather than being equally distribu-tion across the scrotal region and seemsconfined to adult males in breeding condi-tion, this would indicate a probable socialcommunication function, rather than relatedto temperature regulation or a radiationshield.

Discussion

Given the putative association mentionedabove between Ravenala and Myzopoda, we



examine this relationship in finer detail tounderstand potential speciation mechanismsin Myzopoda. Blanc et al. (2003) indicatedthat the upper elevational limit of Ravenala issomewhere between 1000 and 1100m. Theresults of elevational transects of plantcommunities in the eastern humid forestsare concordant with this assessment. Rave-nala, as a member of non-disturbed forest,drops out between 800 and 1000m, and thispattern is relatively consistent across the 121latitudinal range of the eastern forests (Lewiset al. 1996; Messmer et al. 2000). However,Ravenala can be found in isolated patches,particularly in wet valley bottoms associatedwith degraded habitats (savoka), up to about1300m (Humbert 1965). This plant is com-mon along the complete length of the easternlowland area, with a distinctly more clumpeddistribution in drier zones at the northern endof the island near Antsiranana, where it canbe found in wet valley bottoms, and thenwest to the Sambirano region near Ambanja,and then becomes distinctly less common


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Table 4. Factor loadings of principal componentanalysis for a series of analyses of Myzopoda spp.Graphic presentations of these comparisons are givenin Fig. 6.

Variable Factor 1 Factor 2

External measurementsTotal length 0.748 �0.125Tail length 0.475 �0.676Hind foot length 0.261 0.850Ear length 0.824 0.051Forearm length 0.855 0.145Eigenvalues 2.27 1.22% total variance 45.3 24.4

Wing bone measurements3rd digit – metacarpal 0.926 0.2473rd digit – 1st phalanx 0.438 0.6073rd digit – 2nd phalanx 0.219 0.8333rd digit – 3rd phalanx 0.646 �0.0714th digit – metacarpal 0.917 0.2434th digit – 1st phalanx 0.551 0.4744th digit – 2nd phalanx �0.127 0.8535th digit – metacarpal 0.920 0.0435th digit – 1st phalanx 0.280 0.7025th digit – 2nd phalanx 0.046 0.866Eigenvalues 3.60 3.32% total variance 36.0 33.2

Cranial measurementsOccipitonasal length 0.950 �0.125Greatest zygomatic breadth 0.791 0.372Interorbital breadth 0.067 0.987Mastoid width 0.775 0.472Lacrimal width 0.842 0.203Eigenvalues 2.84 1.39% total variance 56.9 27.8

Dental measurementsAnterior palatal width 0.533 0.429Maxillary toothrow �0.046 0.906Upper molariform toothrow 0.396 0.510Width M3 0.908 �0.048Eigenvalues 1.28 1.27% total variance 31.7 31.7


along the west coast south to at leastNamoroka. On the basis of our joint batinventory data in humid forests of the east,using direct capture and acoustic sampling,the highest elevation Myzopoda has beenrecorded is at around 970m, in the vicinity ofAndasibe. Thus, there is considerable con-cordance between the upper altitudinal limits



in the eastern biome of Ravenala andM. aurita.Given the considerable morphological simi-larity between M. aurita and M. schliemanni,and the fact they are the only members of thisendemic genus, it is assumed that one of thetwo represent a speciation event after dis-persal. Further, given their apparent propen-sity for roosting in Ravenala, which is notablymore common in the east than the west, weassume that the species group originated inthe east and then dispersed to the west.(However, a priori the dispersal event andsubsequent speciation could have been in theopposite direction.) Here, we propose twodifferent scenarios on how this event mayhave taken place.The first hypothesis is dispersal directlyacross the Central Highlands, for example,at the latitudinal level of Antananarivo(Fig. 1). This seems unlikely given that thiszone rises notably higher than 1000m and theeast-west hiatus of several hundred kilo-meters for even moderate size patches ofRavenala habitat. However, one cannot ruleout a hypothesis of cataclysmic dispersal,such as cyclone winds that physically blewthe animals across the Central Highlands.The second hypothesis is an east to westdispersal event across the region fallingbetween the Central Highlands and NorthernHighlands, which comprises a vast zone withpotential corridors of less than 1000m inelevation (Fig. 1). Across the northern por-tion of central Madagascar, in the zonelinking Toamasina and Mananara on theeast coast to Mahajanga on the west coast,Ravenala is patchy in its distribution andhence stepping-stone dispersal across thisregion is plausible.Comparisons between the genera Myzopodaand Thyroptera show a remarkable numberof convergent parallels in certain morpholo-gical adaptations and aspects of reproduc-tion. It would appear that females ofT. tricolor are polyoestrous, with perhapstwo breeding cycles per year, and that adultmales have multiple spermatogenic roundsor produce sperm continuously (Wimsattand Enders 1980; Krutzsch 2000). Further,T. tricolor is known to have a complex socialsystem (Vonhof et al. 2004). Members of this


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genus also have relatively complex geogra-phical ranges (Gregorin et al. 2006). Given allof the interesting repeated convergent pat-terns between Myzopoda and Thyroptera,research on the natural history and repro-ductive system in the former genus is almostcertain to uncover some fascinating details.On the basis of limited information,M. schliemanni lives in portions of the centrallowland west where there are relatively mesicconditions, associated with marshes, rivers ortransitional humid-dry forests. Although twoof the three sites this species is knownfrom are protected areas (Parc Nationald’Ankarafantsika and Parc National deNamoroka), it appears, based on our inven-tory work, to be sparsely dispersed in thecentral western portions of the island (Good-man et al. 2005). The forest habitat at theAnkaboka sites, as well as one of the twosites at Namoroka, were notably disturbed byhuman activities and by extrapolation thisspecies appears to be resilient to certain levelsof anthropogenic perturbation. Across itsknown range, measuring about 300 km inlength, there is considerable non-continuoushabitat similar to sites in which it has beenfound, and we suspect that the threats to thisspecies’ long-term existence are not notablysevere.In their assessment of the conservation statusof the world’s Microchiroptera, Hutson et al.(2001) considered M. aurita to be a species ofspecial concern, largely associated with itbeing the sole member of an endemic familyand that little information was availableabout its distribution. Over the course ofthe past decades, largely as a result ofbiological inventories, we now know thatthis species is more broadly distributed thanpreviously realized and inhabits areas ofextensive secondary habitats of Ravenala.While it is true that M. aurita is currentlyonly known from in or around three pro-tected areas (Parc National de Marojejy, ParcNational de Masoala and Reserve Specialed’Andasibe; Pont and Armstrong 1990;Russ et al. 2001; Schliemann and Goodman2003; Tab. 1), inferred levels of threat maybe less severe then previously assumedgiven its ability to live in highly disturbedareas. Finally, based on the description of a



second species of Myzopoda herein, thefamily Myzopodidae is no longer monospe-cific and the sole family of bats withthis level of uniqueness in the world is theCraseonycteridae.The discovery of M. schliemanni and newinformation on the distribution and conser-vation status of M. aurita are based on recentsurvey work. In a few years considerableprogress has been made on numerous aspectsof the chiropteran fauna of the island. Thisunderlines the importance of basic scientificresearch for establishing the priorities forconservation programs and assessments ofpresumed rare and possibly endangeredanimals.

Acknowledgements

We are grateful to the Direction des Eaux etForets and Association Nationale pour laGestion des Aires protegees for issuingcollection and exportation permits. BP Con-servation Programme, Darwin Initiative(162/10/024), John D. and Catherine T.MacArthur Foundation, Rufford SmallGrants and Volkswagen Foundation pro-vided funding for portions of the fieldwork.For access to specimens under their care, weare indebted to Nancy Simmons (AMNH),Paulina Jenkins (BMNH), Jean-Marc Ponsand Ronan Kirsch (MNHN), Chris Smeenk(RMNH), Olga Ramilijaona and DanielRakotondravony (UADBA), Berthe Rako-tosamimanana (UADP), Michael Carleton(USNM), Robert Asher (ZMB) and HaraldSchliemann (ZMH). We are grateful to FanjaRatrimomanarivo and Angelos Tianarifidyfor providing some additional specimenmaterial for this study and Julie Razafima-nahaka, Daudet Andriafidison, and RoselineRampilamanana for aid with fieldwork inNamoroka. Several friends and colleaguesprepared or made available figures: RebeccaKramer (Figs. 4 and 5), Achille P. Raselima-nana (Fig. 3b), John Weinstein (Fig. 2), andLucienne Wilme (Fig. 1). Jorg Ganzhornkindly translated the summary into German.For comments on an earlier version of thisstudy we are grateful to Jon Russ andRichard Jenkins.


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Zusammenfassung

Beschreibung einer neuen Art von Myzopoda (Myzopodidae: Chiroptera) von West-Madagaskar

Die Myzopodidae, eine endemische Familie von Fledermausen auf Madagaskar, galt bisher alsmonospezifisch. In Madagaskar wurde eine neue Art, M. schliemanni, gefunden. Diese Art aus denTrockenwaldern West-Madagaskars unterscheidet sich in der Fellfarbung, externen morphologischenMaXen und Schadelmerkmalen von M. aurita, der bisher einzig beschriebenen Art der Familie.Myzopoda aurita ist aus den immergrunen Regenwaldern des Ostens Madagaskars bekannt.Biogeographische Muster der Verteilung von Myzopoda und ihre enge Assoziation mit dem ‘‘Baumdes Reisenden (Ravenala madagascariensis; Strelitziaceae)’’ werden in Zusammenhang mitSpeziationsmechanismen diskutiert, die zur Entstehung der westlichen und ostlichen Artbeigetragen haben.r 2006 Deutsche Gesellschaft fur Saugetierkunde. Published by Elsevier GmbH. All rights reserved.

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Authors’ addresses:

Steven M. Goodman, Field Museum of NaturalHistory, 1400 South Lake Shore Drive, Chicago, IL60605, USA and WWF, BP 738, Antananarivo (101),Madagascar Felix Rakotondraparany, Departementde Biologie Animale, Faculte des Sciences, Universited’Antananarivo, B.P. 906, Antananarivo (101), Ma-dagascar Amyot Kofoky, Madagasikara Voakajy,B.P. 5181, Antananarivo (101), Madagascar(e-mail: [email protected] (F. Rakoton-draparany), [email protected] (A. Kofoky).


. biol. (2006), doi:10.1016/j.mambio.2006.08.001

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the description of a new species of myzopoda (myzopodidae: chiroptera) from western madagascar

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