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A reassessment of the taxonomic status and
distribution of the subspecies of Lyroderma lyra
(Chiroptera: Megadermatidae)
Lei Feng* |
Northeast Normal University China
Hunan Academy of Forestry China
Dorottya y* |
Department of Zoolog Hungarian Natural History Museumt Hungary
Hungarian University of Agriculture and Life Sciencesödöll Hungary
Roberto Portela Miguez
Department of Life Science Natural History Museum Londo UK
Phaedra Kokkini
Department of Life Science Natural History Museum Londo UK
|
Department of Zoolog Hungarian Natural History Museumt Hungary
Shahneaz Ali Khan
Chattogram Veterinary and Animal Sciences University Bangladesh
Uttam Saikia
North Eastern Regional Centr Zoological Survey of India Risa Colon Shillon
3 India
Shyamkant S. Talmale
Western Regional Centr Zoological Survey of India Pun4 India
Wenhua Yu |
College of Life Sciency China
* These authors contributed equally to this articl
Shaoying Liu |
Sichuan Academy of Forestry China
Tinglei Jiang |
Northeast Normal University China
jiangtl730@nenu.edu.cn
Gábor a* |
Department of Zoolog Hungarian Natural History Museumt Hungary
; ; ;
;
Abstract
Lyroderma lyra is one of the six currently recognised species of the family Megadermatida
L. lyra wherein the nominate sub
species is predominantly distributed in South Asi and the other subspecie L. is
occurring in southern China and Southeast Asi Based on the congruent results of comparative
morphology of skulls and bacul
including both subspecies and incorporating the most extensive geographic sampling of “e”
and cytb sequence we have elevated the Southeast Asian subspecies to a dis
tinct species statu e
Keywords
Megaderma
Introduction
The false vampire belonging to the fam
ily Megadermatida are restricted to the
tropics and subtropics of the Old World
ilson & Mittermeie
nally by a rounded anterior noselea
well simple and erect poste
rior noselea large oval ears joined above
the forehea forked tragu and well
developed interfemoral membrane but no
bony tail ates & Harriso Wilson &
Mittermeie the premax
illae are essentially absen and the nasals
are greatly reduced; and dentall the lack
istics of the family oisook
Wilson & Mittermeie
Six genera are currently recognise each
containing one extant specie namely
istributed in
Africa) Cardioderma cor ast Africa)
ustralia)Megaderma
outheast Asia)
thongareeae eninsular Thailand)
Lyroderma lyra outh and Southeast
Asia) merican Society of Mammalogists
)4; Simmons & Cirranell4;
Volleth 1; Wilson & Mittermeie
Lyroderma Peter
treated as a subgenus of Megaderma
fro but recent phylogenetic recon
structions using various markers ick
Kanuch Soisook
consequently separated Lyroderma
and Megaderma as full gener Soisook
provided a comprehensive
overview of the taxonomicnomenclatural
history of Megadermatidae; for a short
summary of the change see
Within Lyroderma two subspecies are
currently recognised; the nominal form is
widespread in South Asi from Afghani
stan to Banglades whereas L.
riginally described as but
changed into for gender agree
ment) is found in Southeast Asi from
South China to Peninsular Malaysia ingh
& Sharm3; Soisook Wil
son & Mittermeie
distributional limits are not clea The
Myanmar ormerly Burma) population of
Lyroderma
to the taxon “lyra” by Sinha
that his arguments are debatabl He con
between “lyra” and “e” [ being
Nomenclatural history of the genera Megaderma and Lyroderma
(1810)
Andersen &
Wroughton
(1907)
Ellerman &
Morrison
(1951)
Corbet & Hill
(1992)
Simmons
(2005)
Francis
(2019)
Simmons &
Cirranello
(2024)
Megaderma Eucheira*Megaderma Megaderma
(Lyroderma)
Megaderma
(Lyroderma)
Lyroderma Lyroderma
lyra lyra lyra lyra lyra lyra lyra
lyra lyra lyra lyra lyra lyra
caurina caurina
Megaderma Megaderma
(Megaderma)
Megaderma
(Megaderma)
Megaderma Megaderma
carimatae
natunae
* According to Andersen and Wroughton Eucheira Hodgso Lyroderma
Peter2; howeve Eucheira HodgsoEucheira Westwoo Lepidoptera)
ee Mille
the larger taxon] but noted that “Indian
examples have smaller skulls than those
from Burma” inh
The taxon was described by
Andersen & Wroughton as a new
species in the genus Eucheira based on two
skins and skulls collected from Xiame
Fujian Province and Shanto
Provinc South China ndersen &
Wroughto
guished the new species from E. lyra on
kull length of is between 2
the prenasal notch narrowe
rounded posteriorly; v skull length
between 2 and the prenasal
notch posteriorly wide lyra)
ndersen & Wroughto
3) published a study of bats from
Chin in which he measured peci
mens of E. collected in western
Sichua Chin
in Fujia Investigating if the two popu
lations under study may represent two
distinct form based on the external and
craniodental measurements taken by hi
lations cannot be distinguished neither
Despite the limitations
of his study anbor 3)
ther explanation he listed as a sub
species of L. lyra the view that is widely
accepted by subsequent authors ates &
Harriso Corbet & Hil1; Csorba
& Topá4; Ellerman &
1; Lekagul & McNeel Sanbor
3; Simmon Simmons &
Cirranell 4; We 3; Wilson &
Mittermeie Wilson & Reede
The karyology of the two subspecies
has been studie and they share the
same karyotype u & Harad
Nevertheles the obvious morphological
ummarised by Csorba & Topá 4) have
never been tested by molecular biologi
cal method In this stud we applied an
integrative taxonomic approac
ing analyses of morphological data and
mtDNA sequences based on the most
comprehensive taxonomic samplings to
investigate the taxonomic status of L. lyra
lyra and L.e
Methods
Specimens examined
For morphological comparison and partly
for multivariate statistic
specimens of L. lyra and eleven specimens
of
mo widely distributed and frequently
were use The specimens are housed in
the following collection
of Life Science
Chin
Natural History Museu Budapes Hun
gar
Changsh Huna Chin
east Normal Universit Changchu Jili
Chin
Londo United Kingdo
of Songkla University Zoological Collec
tio Hat Ya Thailan
Academy of Forestr Chengd Sichua
Chin
Natural Histor Ljubljan Sloveni
si Bang Malaysi
Zoological Survey of Indi Pune and
Zoological Survey of Indi Shillon The
distribution map of the sampling points is
shown in
Lyroderma lyra lyra
2
sex unknown;
42
n;
n;
2
e
1 1
sex
unknown;
Lyroderma lyra riangles)
false vampire bat quares) and samples analysed in the present pape Full
symbol both morphological and genetic data were used; empty symbol only molecular data
were available; symbol only morphological information was use Some symbols
represent multiple specimens from the same localit
2 not
registere sex unknown;
1
sex unknow
Lyroderma lyra ss
2
42 2
2 2
3
a
sex unknow
1
Measurements and morphometric
analysis
We took one external orearm length) and
13 craniodental measurement following
Bates and Harrison
shown in The external measure
the cranial
and dental ones were measured under a
stereomicroscope by digital vernier calli
per to the nearest Measurements
were recorded from adult specimens onl
Previous studies have indicated that
the forearm of female individuals of L. l.
lyra is longer than that of male individuals
inh
dent samples t show that the fore
arm length of female individuals of L. l.
male individuals T2 2)
Since the craniodental parameters did
not meet the homogeneity of variance
we employed the U test to
between male and female individuals of
L. l. and L. l. lyra acha
The skull length ) and width ) of
female L. l.
than those of male L. l. e hence
fort the analysis of craniodental param
eters was conducted separately for the
two sexe We employed raniodental
measurements of pecimens subset
of the morphologically studied museum
materials) ndividuals each for
females and males and 4)
form principal component analysis )
and linear discriminant analysis )
distributions or outlier we performed
logarithmic transformation on the data
before conducting and analyse
The results showed that the and
outcomes before and after logarithmic
transformation were simila The analy
ses were carried out using R Core
Tea 4)
the Whitney U Tes
of the results was accomplished using the
ggor ggplot plot and factoextra pack
ages in R Core Tea 4)
assess the suitability of the data for use
Description of crania mandibula and external dimensions and their abbreviations
Measurements Acronyms Descriptions
Forearm length From the elbow to the distal end of the forearm including
carpals
of skull
From the apex of the upper internal incisors to the occiput
length
From the anterior surface of the upper canines to a line
connecting the occipital condyles
Interorbital
breadth
Least breadth between the orbits
Zygomatic width
arches
Mastoid width
Braincase width
Braincase height Braincase height posteriorly to the auditory bullae
Maxillary canine
outer width
Distance between outer margins of C1
Width of upper
molars
M3M3W Maximum distance between the bases of the outer edges of
the third posterior molars on each maxillary side
Maxillary
toothrow length
Maximum length from the anterior margin of the maxillary
canine to the posterior margin of the base of the last molar on
the same side
Mandible length The length from the most anterior edge of the mandible
xcluding incisors) to the most posterior edge of the
mandible
Mandibular
toothrow length
cm3L Maximum length from the anterior edge of the mandibular
Mandibular
molariform
toothrow length
m1m3L Distance from the anterior edge of the m1 alveolus to
the posterior edge of the m3 alveolus of the mandibular
posterior molar
in
) test and the Bartlett’s test of sphe
ricity focusing on the skull datase The
Olkin measure of sampling
adequacy ) was greater than
and the Bartlett’s test of sphericity was
1)
data was suitable for askey
age value
Baculum
Preparation and preservation of the penial
bone followed that of Friley
Selected measurements of male group L.e L. lyra and M.a and vector correlation loadings with original variables of principal
components and ) and discriminant functions and ) for selected samples of male group L. l.e L. l. lyra and M.a See
Table 2 for variable abbreviation
Meareplsurements L. l. sinense L. l. lyra M. spasma p-value
M3M3W
cm3L
m1m3L
Selected measurements of female group L.e L. lyra and M.a and vector correlation loadings with original variables of principal
components and ) and discriminant functions and ) for selected samples of female group L. l.e, L. l. lyra and M.a See
Table 2 for variable abbreviation
Measurements L. l. sinense L. l. lyra M. spasma p-value
baculum was draw and its maximum
length was measured using a Mitutoyo dig
ital calliper to the nearest m under a
stereomicroscop
Phylogenetic inference
Total genomic was extracted with
DNeasy Blood & Tissue Kit iage
y) according to the instructions of
the manufacturer from
Lyroderma specimens
forming part of the genetic resources’ col
lections of
Pune and Shillon
For phylogenetic analysi the mitochon
drial cytochrome oxidase c subunit
and the cytochrome b gene cytb
tyrLA assanin 2)
and báñ
Weyeneth
reactions were performed in µl using
1 µl of genomic µl of the
primers µl of
wate µl of dNT µl of DreamTaq
USA)
and µl of DreamTaq Polymerase
USA)
works were done for respective
specimens housed in a particular museu
The and cytb sequences were aligned
of Staden Package
The newly generated sequences were com
pared with 43 ncluding utgroups)
and 14 cytb ncluding utgroups) gene
sequences of false vampires downloaded
from the National Center for Biotechnology
Information ) supple
mentary table S3)
The phylogenetic analysis was per
formed using the maximum likelihood
) metho in the software
umar
por we computed a nonparametric boot
Additionall
the genetic distance of
and cytb genes was calculated by Bootstrap
method using
Results
Morphology and multivariate statistics
We could not establish any statistically
aired samples
T F 3 between L. l.
and L. l. lyra
with respect to forearm
length )
The U Test was used to
compare thirteen cranial parameters of
males and females between two subspe
cie
L. l.
lyra and L. l.e On averag within the
same se the skull of L. l. is longer
and wider than that of L. l. lyra
and supplementary 1 and S2)
despite that overlapping values in all mea
surements were observe Howeve in
line with the observations of ndersen
& Wroughto
interorbital region and the prenasal notch
taxa ee details in the Introduction and
in
)
% of
the total craniometric variation and
the skull A and 4A)
axi there is no overlap between the
groups of male M. and L. l. lyra
or between M. and L. l. e;
these groups show some overlap along
between M. and L. l. lyra
or between M. and L. l.
male L. l. lyra and L. l.
exhibit overlap along both and
Along the axi there is no over
lap among female M.a L. l. lyra and
L. l.e; howeve on the axi all
three groups show overlap
both male and female group the fac
tor loadings show that the loadings for
are very clos while on the
second the factor loadings for
and M3M3W are higher than those of
the other sample indicating that they
are the most useful discriminant factors;
following thes the next important dis
criminant factors include and
and B and 4B)
factor loadings of other
supplementary tables S1 and S2 The
of craniodental measurements shows
that in there is no overlap among the
The dorsal and ventral views of the skulls of L.
) from Bac Tha Vietnam c)
L. lyra ) from West Benga India d)
e;t;
notch;e;
three species for both males ) and
females there is
some overlap among the three species in
male and only between L. l. and
M. in females C and 4C)
male some parameters associated with
the length or width of the skull and max
illa ) are most help
ful in distinguishing these three groups
table 3) howeve
some parameters related to the width of
the skull and maxilla M3M3
)
these three groups table 4)
The baculum of both L. l. and
L. l. lyra consists of two separat
bones only partially connected with liga
ments; the penis bones of L. l. sinense are
distinctly longer than those of L. l. lyra
m v m) and straight
distinctly bent dorsoventrally in L. l.
lyra)
Phylogenetic reconstructions
The newly obtained sequences included
and cytb sequences for eight L. l.e
L. l. lyra and one M.a All novel
sequences were deposited in the
and vector correlation B) and and vector
correlation D) of cranial measurements for male reen dots)
Lyroderma lyra ed dots) and lue dots)
and vector correlation B) and and vector
correlation D) of cranial measurements for female reen dots)
Lyroderma lyra ed dots) and lue dots)
The dorsal eft) and lateral ight) views of the baculum of a) L.
) from Bac Tha Vietnam; and L. lyra )
from Maharasthr Indi1 mm
Phylogenetic tree resulting from the Maximum Likelihood analysis of
sequences of species of Megadermatida numbers indicate nodal
suppor the outgroup genes were derived from species of the genera
and
Phylogenetic tree resulting from the Maximum Likelihood analysis of cytb sequence of species
of Megadermatida numbers indicate nodal suppor the outgroup genes were derived from
species of the genera and
supplementary table S3
The phylogenetic tree constructed
based on and Cytb sequences revealed
similar phylogenetic relationship with
L. l. being closely related to and
forming a sister clade with L. l. lyra while
M. grouped with species of other
genera within the Megadermatidae fam
il despite the low nodal support for this
phylogeny and branching events
and supplementary )
between L. l. and L. l.
lyra for is 1 it is
age for the two mitochondrial
genes between Lyroderma and Megaderma
oth are higher than 1; and
Discussion
Although there have been variable
amounts of overlap among all the mea
sured craniodental character we
of L. lyra The results of the of the
nine craniodental measurements showed
slight overlap between L. l. and
L. l. lyra male whereas the results of the
showed no overlap in between
male and female L. l.e L. l. lyra and
M. a
The morphological diversity of the
mammalian baculum is subject to direct
selection as a result of their role in copu
lation
Stockle 2)
of the baculum of adult individuals is
Average within Megadermatidae based on sequence Boldface value indicates
the distance between and Lyroderma lyra
Taxa 1 2 3 4 5 6 7
Lyroderma lyra
thongareeae
Cardioderma cor
Average genetic distance within Megadermatidae based on cytb sequence
Boldface value indicates the distance between and Lyroderma lyra
Taxa 1 2 3 4 5 6
Lyroderma lyra
Cardioderma cor
considered a diagnostic criterion for spe
ouangboubpha
Latorr 3; Soisook
Srinivasulu Thoma
Vercillo & Ragn 1)
the baculum of a Lyroderma collected
in Thailand was about 1 mm oisook
of an Indian specimen was about m
rinivasulu
our result
which supports that bacular
features play an important role in the clas
oisook
length between the two tax but the simi
larity in structur supports the idea that
L. and L. lyra are two separat but
closely related specie
The phylogenetic reconstructions using
and cytb gene sequence conse
quently placed all Lyroderma samples into
a single clad sister to a clade containing
all other gener This result is consistent
with other recent phylogenies and sup
ports the validity of Lyroderma as a sepa
rate genus ick Kanuch
Soisook
Phylogenetic analyses also evidenced
that L. lyra is divided into two independent
clade and the genetic distances between
the and cytb gene fragments were
more than 12% and nearly 1 between L.
lyra lyra and L.e respectivel In
line with the widely accepted view radle
& Bake 1) this magnitude of genetic
statu The congruent results of the mor
phologica morphometri and molecular
analyses in the present study strongly sup
port the view of Andersen and Wroughton
that the genus Lyroderma consists
of two specie L. lyra and L.e
The critical evaluation of previous
work based on craniodental measure
ments and the structure of the interorbital
regio helped us to clarify the taxonomic
status of the populations in Afghanistan
and Pakistan where neither directly com
parable museum specimens nor molecu
lar data were available fro
reported the species from Jhelu
Punja Pakista and as it can be judged
from the quality photograph on
the bases of the shape of the interorbital
regio anterior narial emargination and
the interpterygoid region
lation is more likely to represents L. lyra
provided measurements he
trait which proved to be a reliable distin
guishing character between “lyra” and
“e” and table 3) of three
Afghani specimens ange m)
Both above publications concur with
the view of previous authors ates
& Harriso Corbet & Hil 2;
Khajuri Srinivasulu & Srinivasul
2)
of Lyroderma belongs to L. lyra
The Myanmar specimens held in the
collection of robably these are
the same individuals as listed by Sinha)
“lyra” given by
Sinha ee the upper mensural
extremes of L. lyra given in table 3)
the morphology of the interorbital regio
narial emarginatio and the interptery
goid regio indee
specimens as L. lyra
Conclusion
included both former subspecies of L. lyra
in a phylogenetic framework and used the
widest geographic sampling of the taxon
“e” so fa
ment
the taxa “lyra” and “e”
were corroborated by multivariate statis
tical analyses and ) and by phy
logenetic tree reconstructions based on
mitochondrial marker This integrative
approach provides solid evidence that the
two former subspecies represent separate
specie L. lyra and L.e Additionall
our results also supported that the west
ernmost and Myanmar populations of
Lyroderma belongs to L. lyra To sum u
L. is distributed in southern Chin
Thailan Lao Vietna Cambodi and
Malaysi whereas L. lyra occurs from
Afghanistan to Myanmar
presence of L. lyra can be assumed in east
Myanma future studies should involve
the collection of more samples from that
country and from the surrounding area
Myanmar populations and to determine
the precise distribution boundaries of the
two specie
Acknowledgements
Li Xin and Li Xinyao are acknowledged for
their invaluable suggestions on an earlier
version of the manuscrip The prepara
tion of the geographical distribution map
was greatly enhanced by the contributions
of Li Dianjun and Wang Zhiqian Dai
Wentao and Shi Shengchao are thanked
for their assistance in specimen collectio
Furthermor Liu Yingxun is appreciated
for her help with morphometric analysi
Boris Krystufek )
) )
kindly provided access to the specimens
under their car We thank three anony
Author contributions
Fen methodolog
formal analysi
n; röss data curatio
methodolog formal analysi
review & editing; Portela Mique
resource g;
Kokkin resource
editing;örfö
in supervisio funding acquisition; Ali
Kha resource funding acquisition;
Saiki resource
editing; Talmal resources; Y resources;
Li resources; Jian
editin supervisio funding acquisition;
Csorb resource
supervisio
funding acquisitio
to declar
Funding
This research received support from the
National Natural Science Foundation of
China rant 2) and the Spe
cial Foundation for National Science and
Technology Basic Research Program of
China 1) to TLJ; from the
by the European Community Research
Infrastructure Action under the
“Capacities” Program and by the National
Researc Developmen and Innovation
Fund of Hungary
Supplementary material
Supplementary material is available online
a
httpor
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