BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions,research libraries, and research funders in the common goal of maximizing access to critical research.

Postnatal Growth and Age Estimation in Marshall's Horseshoe Bat,Rhinolophus marshalliAuthor(s): Longru Jin, Luo Bo, Keping Sun, Ying Liu, Jennifer Pan Ho and Jiang FengSource: Acta Chiropterologica, 14(1):105-110. 2012.Published By: Museum and Institute of Zoology, Polish Academy of SciencesDOI: http://dx.doi.org/10.3161/150811012X654312URL: http://www.bioone.org/doi/full/10.3161/150811012X654312

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological,and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and bookspublished by nonprofit societies, associations, museums, institutions, and presses.

Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance ofBioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.

Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercialinquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

INTRODUCTION

Most newborn bats are altricial and experience

a postnatal growth period in which they develop

appropriate sensory and locomotor skills to become

independent from their mothers (Baptista et al.,2000). Many researchers have studied the changes

in behavior, physiology, and ecology of this period

to investigate life history traits and to estimate ma-

ternal investment (Kunz et al., 2009).

There have been numerous studies on postnatal

growth of bats under both natural (Hoying and

Kunz, 1998; McLean and Speakman, 2000; Chaver -

ri and Kunz, 2006; Allen et al., 2009; Liu et al.,2009a; Jin et al., 2010) and captive conditions

(Boyd and Myhill, 1987; Rajan and Marimuthu,

1999; Elangovan et al., 2003, 2007; Raghuram and

Marimuthu, 2007). It has been shown that measure-

ments of body mass, forearm length and total length

of the epiphyseal gap of the fourth metacarpal-pha-

langeal joint (De Fanis and Jones, 1995; Hoying and

Kunz, 1998) can be used to estimate the age of bats.

Forearm length is the most accurately measured and

the most reliable characteristic for estimating age

during the early linear growth period of bats. How -

ever, the length of the epiphyseal gap is best for

estimating age at later stages of postnatal growth

(Kunz and Anthony, 1982; Krochmal and Sparks,

2007). Body mass is less reliable for estimating the

age of growing bats because it is highly sensitive to

variations in short term nutritional intake, energy

expenditure, and daily water flux (Stern and Kunz,

1998).

Accurate age determination is important for be-

havioral, physiological and ecological studies (Kunz

and Hood, 2000). In the absence of age estimates, it

is impossible to determine growth rates, the timing

of sexual maturity, the periodicity of reproduction,

the development of various behavioral repertoires,

or the longevity of an animal (Elangovan et al.,2003). In addition, patterns of growth and develop-

ment vary among species and families of bats.

Growth parameters derived from nonlinear models

(e.g., logistic, Gompertz, and von Bertalanffy) are

especially valuable for drawing interspecific com-

parisons because growth equations are independ-

ent of body size and duration of the postnatal

growth period (Kunz and Robson, 1995). However,

Acta Chiropterologica, 14(1): 105–110, 2012PL ISSN 1508-1109 © Museum and Institute of Zoology PAS

doi: 10.3161/150811012X654312

Postnatal growth and age estimation in Marshall’s horseshoe bat,

Rhinolophus marshalli

LONGRU JIN1, 2, LUO BO1, 2, KEPING SUN1, 2, YING LIU1, 2, JENNIFER PAN HO3, and JIANG FENG1, 2, 4

1Jilin Key Laboratory of Animal Resource Conservation and Utilization, Northeast Normal University, Changchun 130024, China2Key Laboratory for Wetland Ecology and Vegetation Restoration of National Environmental Protection,

Northeast Normal University, Changchun 130024, China3Department of Integrative Biology and Physiology, University of California, Los Angeles, USA

4Corresponding author: E-mail: fengj@nenu.edu.cn

Based on mark-recapture data, we studied the postnatal growth of Marshall’s horseshoe bat (Rhinolophus marshalli) in Hekou

County, Yunnan Province, China. Our results detected no significant differences both in body mass and in forearm length between

males and females at birth. On average, young bats were not agile fliers until 31 days of age. Body mass and forearm length of pups

followed a linear pattern of growth until day 13, with mean growth rates of 0.17 g/day and 1.22 mm/day, respectively, and thereafter

growth rates decreased. Length of the total epiphyseal gap of the fourth metacarpal-phalangeal joint showed a linear increase up to

10 days followed by a linear decrease until day 40 with a mean rate of 0.09 mm/day. Together, two equations permitted estimation

of the age of R. marshalli pups between 1 and 40 days of age. Of the three nonlinear growth models (logistic, Gompertz, and von

Bertalanffy), the logistic equation provided the best fit to the empirical curves for body mass and forearm length.

Key words: Rhinolophus marshalli, postnatal growth, age estimation, nonlinear growth models

comparative studies among different taxa should be

based on the same model (Zullinger et al., 1984).

Most studies on postnatal growth in bats were

conducted on vespertilionids in temperate zones,

but only a few focused on rhinolophid bats (Sharifi,

2004a; Dietz et al., 2007; Funakoshi et al., 2010).

Marshall’s horseshoe bat (Rhinolophus marshalli) is a widespread species, ranging through Thailand,

Bur ma, Vietnam, Laos Peninsular Malaysia, and

more recently, China (Simmons, 2005; Zhang et al.,2009). However, little information is available on its

postnatal growth. Thus, our aims were to measure

the sizes of bat pups at birth, to derive age estima-

tion equations based on forearm length and the

length of the total epiphyseal gap, and to compare

the growth patterns of body mass and forearm length

using three non-linear equations.

MATERIALS AND METHODS

This study was conducted in Banshan Cave (22°36’N,

103°50’E, ca. 80 m long, 1.8 m wide, and 2 m high) in Hekou

County, Yunnan Province, China. The cave housed a mixed

colony of ca. 70 R. marshalli and 400 Hipposideros pomona.

The surrounding area was covered by dense forest of mostly

rubber tree (Hevea brasiliensis).

Prior investigation indicated that female R. marshalli un-

dergo parturition from late April to early May. Therefore, we

checked the cave daily from about 15 days before parturition

throughout the season (a total of 18 days). We hand-captured

neonates immediately following the nightly emergence of

adults. Neonates with an attached umbilical cord were assumed

to be 1 day old (Kunz and Robson, 1995). After the sex of each

pup was determined, a numbered aluminum alloy ring (Porzana

Ltd., United Kingdom) was placed on the forearm of each bat

for individual identification. In total, 31 neonates with umbilical

cords were marked.

Body mass was recorded to the nearest 0.01 g using an elec-

tronic balance (ProScale LC-50, United States). Forearm length

was measured to the nearest 0.01 mm with digital vernier

calipers (TESA-CAL IP67, Switzerland). The total length of the

epiphyseal gap was measured to the nearest 0.01 mm using

calipers while the wing of the bat was spread over a transparent

solid plastic sheet illuminated from below with a strong flash-

light to visualize the gap (Sharifi, 2004b). To minimize errors in

using vernier calipers, individual measurements were repeated

three times and the means were used in the analysis.

To limit disturbance and possible abandonment, marking of

young and measuring morphological characters of pups took

place when adults were away in the foraging areas and the pro-

cedure was completed within 1.5 hours. As soon as all young

had been measured and weighed, they were returned to

their sites, as close to their original locations in the cave as pos-

sible, before any females returned. The cave was visited every

three days (Table 1) and continued until the majority of pups

were volant and could no longer be captured, even with the aid

of mist-nets and a hand-net by the middle of June 2010.

Capturing of bats was performed with permission from the local

government.

An independent sample t-test was used to compare forearm

length and body mass of males and females at birth. Linear re-

gression equations were derived to predict age from pooled data

for forearm length (1–13 days) and total epiphyseal gap (10–40

days). To derive an age-predictive equation from the measure-

ments, the axes on the growth curve were reversed and age for

the specific periods were considered as the dependent variable

(Kunz and Anthony, 1982). Ninety-five percent confidence and

prediction intervals were plotted for the regression equations for

forearm length and total gap. In addition, growth data of body

mass and forearm length in young bats were fitted to the

three models: logistic, Gompertz and von Bertalanffy equations

(Zul linger et al., 1984). The equations were as follows:

logistic: W = A (1 + exp (-K (t - I)))-1

Gompertz: W = A exp (-exp (-K (t - I)))

von Bertalanffy: W = A (1 -(1 / 3) exp (-K (t - I)))3

where: A is the asymptotic value (g), W is the body mass (g) at

age t (days), K is the growth rate constant (day-1), and I is the

age at the inflection point (days). The parameters A and K in

each model were estimated for the growth of mass in the

neonate population. Similar equations were used for forearm

length. The Levenberg-Marquardt algorithm was used to derive

the best fit to the three nonlinear equations. Results from the

three models were compared by the goodness of fit obtained

from each model (Zullinger et al., 1984). All statistical analyses

were conducted using SPSS ver. 15.0 (SPSS Inc., Chicago, IL,

USA) and data were described using mean ± SD unless stated

otherwise.

RESULTS

Each adult observed females produced a single

offspring. For 31 newborns with umbilical cords, no

significant difference was found in forearm length

between males and females (t = -1.16, d.f. = 29,

P > 0.05). Similarly, body mass of male and female

did not differ significantly (t = -0.78, d.f. = 29,

P > 0.05). The mean values of forearm length and

body mass of newborns were 17.05 ± 0.41 mm and

2.85 ± 0.14 g, and they were 37.4% and 36.3% of

that of adult females, respectively.

At birth, young R. marshalli were naked and pink

with closed eyes, folded ears, and deciduous teeth.

106 L. Jin, L. Bo, K. Sun, Y. Liu, and J. Pan Ho

TABLE 1. Number of individuals captured in Banshan Cave

ParameterDay of sampling

1 4 7 10 13 16 19 22 25 28 31 34 37 40

Number of re-captured bats 31 28 26 27 24 25 23 21 18 19 15 13 8 6

Marked bats re-captured (%) – 90 84 87 77 81 74 68 59 61 48 42 26 19

Seven days after birth, the short, fine, and soft hair

of the pups was distinguishable. Sixteen days after

birth, their ears were erect and some pups’ eyes were

completely open. After 22 days, some young were

able to flutter and glide when they were released by

hand. After 31 days the mean body mass and fore-

arm length were 80.3% and 96.6% of adult values,

respectively. Additionally, most young bats were

observed to fly freely with gentle turns in the cave.

Empirical growth curves of changes in body

mass, forearm length, and length of the total gap of

the fourth metacarpal-phalangeal joint were derived

from the data collected from the 31 young (Fig. 1).

During the first 13 days body mass (Fig. 1A) and

forearm length (Fig. 1B) increased linearly with

growth rates of 0.17 g/day and 1.22 mm/day, respec-

tively. Subsequently, growth rates of these two char-

acteristics decreased to a relatively constant value.

In contrast, the length of the epiphyseal gap in-

creased linearly until day 10 and then decreased

linearly to a mean rate of 0.09 mm/day (Fig. 1C).

Age was highly correlated with forearm length

up to 13 days, by which it grew to 31.72 ± 0.64 mm.

Thus, the age of R. marshalli with forearm length

≤ 31.72 mm could be estimated by the following

equation:

Age (days) = (0.81 × Forearm length) – 12.82

[r2 = 0.99, n = 136, P < 0.01]

Similarly, the age of R. marshalli with forearm

length ≥ 28.01 mm (the mean at day 13) was esti-

mated by the following equation:

Age (days) = (-10.21 × Length of total gap) + 48.36

[r2 = 0.95, n = 199, P < 0.01]

Figs. 2A and 2B show the equations with 95%

confidence and prediction intervals of the estima-

tions of age, using forearm length and length of epi-

physeal gap, respectively. Together, these two equa-

tions allow us to predict the age of young R. mar-shalli from 1 to 40 days after birth. The logistic,

Gom pertz, and von Bertalanffy models gave a good

fit to the changes in body mass and forearm length

(Table 2). Based on the sum of squares model, the

logistic model appeared to provide the best fit for

body mass and forearm length.

DISCUSSION

For newborn R. marshalli, no significant differ-

ences were found in either body mass or forearm

length between males and females, which was con-

sistent with studies on Tadarida brasiliensis mexi-cana (Kunz and Robson, 1995) and Hipposide ros

Postnatal growth and age estimation in Rhinolophus marshalli 107

FIG. 1. Empirical growth curves for (A) body mass, (B) forearm

length, and (C) the length of total gap of the fourth metacarpal-

phalangeal joint of young R. marshalli from day 1 to 40. Some

points represent more than one observation and ‘a’ in (A) and

(B) represents the values of adults

terasensis (Cheng et al., 2002). However, a signifi-

cant difference in body mass was indicated between

sexes of newborn Artibeus watsoni (Chaverri and

Kunz, 2006). In addition, the average size of R. mar-shalli pups at birth (36.3% of female body mass)

was larger than that of most bat species (e.g., Phyl-lo stomus hastatus — 20.7%, Stern and Kunz, 1998;

Rhi nolophus hipposideros — 34.5%, Reiter, 2004;

R. mehelyi — 28.5%, Sharifi, 2004a). Large neonatal

Length

of

epip

hyseal gap (

mm

)F

ore

arm

length

(m

m)

Body m

ass (

g)

Age (days)

Body mass versus age Forearm length versus ageGrowth model Parameter

Estimate SE CV(%) Model sum of squares Estimate SE CV(%) Model sum of squares

LogisticA 6.39 0.044 0.69 10.93 47.38 0.18 0.38 140.09

K 0.11 0.003 2.78 0.10 0.001 0.96

I 3.98 0.159 4.00 6.39 0.091 1.42

Gompertz A 6.59 0.059 0.89 11.28 49.50 0.265 0.54 154.94

K 0.08 0.003 3.75 0.07 0.001 1.37

I -0.16 0.147 -93.63 2.02 0.078 3.87

von Bertalanffy A 6.69 0.068 1.02 11.53 50.63 0.322 0.64 168.18

K 0.07 0.002 2.82 0.06 0.001 1.59

I -2.16 0.166 -7.68 -0.22 0.076 -34.23

Note: A — asymptotic value of body mass (g) or forearm length (mm); K — growth rate constant; I — inflection point; SE — standard error;

CV — coefficient of variation

size is generally associated with an advanced stage

of development and is believed to aid in the conser-

vation of heat generated to maintain constant body

108 L. Jin, L. Bo, K. Sun, Y. Liu, and J. Pan Ho

TABLE 2. Growth parameters in R. marshalli derived from the logistic, Gompertz, and von Bertalanffy growth models

FIG. 2. Equations of age estimation derived from regression line

analysis in R. marshalli, including 95% confidence interval

(narrow band), and 95% prediction interval (wide band) for the

values of forearm length (A) for the first 13 days, and the length

of total gap of the fourth metacarpal-phalangeal joint (B) from

day 10 to 40. Some points represent more than one observation

Length of epiphyseal gap (mm)

Forearm length (mm)

Age (

days)

temperature when the mother leaves the roost to for-

age (Reiter, 2004). Carrying a heavier foetus during

pregnancy and the associated increase in wing load-

ing, however, can reduce flight maneuverability of

pregnant females, which may in turn have increased

predation risks from raptorial birds (McLean and

Speakman, 2000). Liu et al. (2009b) suggested that

R. marshalli are agile fliers due to their low wing

loading, and we concur that carrying a heavier foe-

tus during pregnancy may not have a detrimental an

effect on the mother’s foraging capabilities.

Sustained flight of young bats occurs when they

are 31 days old, by achieving over 80% of adult

body mass and 96% of adult forearm length. This is

similar to juveniles of other insectivorous bats,

which typically begin to fly when they attain 70% of

adult body mass (Barclay, 1995) and 90% of adult

forearm length (Chaverri and Kunz, 2006). Having

a relatively small body mass and large skeletal size

when recently becoming volant lowers wing load-

ing, which in turn increases maneuverability, de-

creases the cost of flight, and thus offers advantages

to young bats that are learning how to detect and

capture flying insects (Hughes et al., 1995).

According to age estimations of R. marshalli,forearm length is useful in determining the age of

the young during the early stage of postnatal growth

when forearm length growth is linear. The length of

the total gap of the fourth metacarpal-phalangeal

joint is useful as an additional parameter for age

estimation during the later stage, when growth of the

forearm becomes non-linear. However, postnatal

growth rates in bats are influenced by climate, food

supply, habitat, latitude, maternal factors, and social

environment (Cumming and Bernard, 1997; Hoying

and Kunz, 1998; Hood et al., 2002; Dietz et al.,2007). Thus, we agree with the suggestion made

by Kunz et al. (2009), to compare the variations in

growth in different years and to generate geographi-

cally specific age equations whenever possible.

Growth curves as empirical models usually have

few parameters and are meant to reflect the nature

and dynamics of the underlying biological process-

es (Karkach, 2006). In our analysis, growth data for

forearm length and body mass were best described

by the logistic equation. The logistic model reflects

the rapid attainment of adult forearm length and

body mass, is computationally simple, and has bio-

logical relevance (Kunz and Robson, 1995). Similar

conclusions were made for Pipistrellus pipistrellus(Hughes et al., 1995), Plecotus auritus (De Fanis

and Jones, 1995) and T. b. mexicana (Kunz and

Robson, 1995). Given the relatively few studies that

have compared different models, the general signif-

icance of these differences remains unclear. In ad-

dition, on the basis of reviews by Kunz and Hood

(2000), growth constant rates of body mass of all

tropical microchiropterans were 0.04–0.11 (n = 11)

and the rates of most temperate microchiropteran

species were 0.12–0.25 (n = 13). In our study, the

growth constant rate of body mass was 0.10, which

embodied the regional features, i.e., a transition zone

between subtropical and tropical environments.

ACKNOWLEDGEMENTS

We thank Zheng Liu and his family for their invaluable field

assistance. This study was financed by the National Natural

Science Foundation of China (Grant Nos. 31030011, 31100305,

and 30900132) and China Postdoctoral Science Foundation

(Grant Nos. 20100481044 and 201104520).

LITERATURE CITED

ALLEN, L. C., C. S. RICHARDSON, G. F. MCCRACKEN, and T. H.

KUNZ. 2009. Birth size and postnatal growth in cave- and

bridge-roosting Brazilian free-tailed bats. Journal of Zool -

ogy (London), 280: 8–16.

BAPTISTA, T. L., C. S. RICHARDSON, and T. H. KUNZ. 2000.

Postnatal growth and age estimation in free-ranging bats:

a comparison of longitudinal and cross-sectional sampling

methods. Journal of Mammalogy, 81: 709–718.

BARCLAY, R. M. R. 1995. Does energy or calcium availability

constrain reproduction by bats? Symposia of the Zoological

Society of London, 67: 245–258.

BOYD, I. L., and D. G. MYHILL. 1987. Variations in the post-

natal growth of pipistrelle bats (Pipistrellus pipistrellus).

Jour nal of Zoology (London), 213: 750–755.

CHAVERRI, G., and T. H. KUNZ. 2006. Reproductive biology

and postnatal development in the tent-making bat Artibeuswatsoni (Chiroptera: Phyllostomidae). Journal of Zoology

(Lon d on), 270: 650–656.

CHENG, H. C, L. L. LEE, and W. L. GANNON. 2002. Postnatal

growth, age estimation, and sexual maturity in the For mo -

san leaf-nosed bat (Hipposideros terasensis). Journal of

Mam malogy, 83: 785–793.

CUMMING, G. S., and R. T. F. BERNARD. 1997. Rainfall, food

abundance and timing of parturition in African bats. Oecol -

ogia, 111: 309–317.

DE FANIS, E., and G. JONES. 1995. Postnatal growth, mother-

infant interactions and development of vocalizations in the

vespertilionid bat Plecotus auritus. Journal of Zoology

(Lon don), 235: 85–97.

DIETZ, C., I. DIETZ, and B. M. SIEMERS. 2007. Growth of horse-

shoe bats (Chiroptera: Rhinolophidae) in temperate conti-

nental conditions and the influence of climate. Mammalian

Biology, 72: 129–144.

ELANGOVAN, V., E. Y. S. PRIYA, H. RAGHURAM and G. MARI -

MUTHU. 2003. Postnatal development in the Indian short-

nosed fruit bat Cynopterus sphinx: growth rate and age esti-

mation. Acta Chiropterologica, 5: 107–116.

ELANGOVAN, V., E. Y. S. PRIYA, H. RAGHURAM and G. MARI -

MUTHU. 2007. Wing morphology and flight development in

the short-nosed fruit bat Cynopterus sphinx. Zoology, 110:

189–196.

FUNAKOSHI, K., E. NOMURA, M. MATSUKUBO, and Y. WAKITA.

2010. Postnatal growth and vocalization development of the

lesser horseshoe bat, Rhinolophus cornutus, in the Kyushu

district, Japan. Mammal Study, 35: 65–78.

HOOD, W. R., J. BLOSS, and T. H. KUNZ. 2002. Intrinsic and ex-

trinsic sources of variation in size at birth and rates of post-

natal growth in the big brown bat Eptesicus fuscus (Chiro -

ptera: Vespertilionidae). Journal of Zoology (London), 258:

355–363.

HOYING, K. M., and T. H. KUNZ. 1998. Variation in size at birth

and postnatal growth in the insectivorous bat Pipistrellussubflavus (Chiroptera: Vespertilionidae). Journal of Zoology

(London), 245: 15–27.

HUGHES, P. M., J. M. V. RAYNER, and G. JONES. 1995. Ontogeny

of ‘true’ flight and other aspects of growth in the bat Pi -pistrellus pipistrellus. Journal of Zoology (London), 236:

291–318.

JIN, L. R., A. Q. LIN, K. P. SUN, Y. LIU, and J. FENG. 2010.

Postnatal growth and age estimation in the ashy leaf-nosed

bat, Hipposideros cineraceus. Acta Chiropterologica, 12:

155–160.

KARKACH, A. 2006. Trajectories and models of individual

growth. Demographic Research, 15: 347–400.

KROCHMAL, A. R., and D. W. SPARKS. 2007. Timing of birth

and estimation of age of juvenile Myotis septentrionalis and

My otis lucifugus in west-central Indiana. Journal of Mam -

mal ogy, 88: 649–656.

KUNZ, T. H., and E. L. P. ANTHONY. 1982. Age estimation and

post natal growth in the bat Myotis lucifugus. Journal of

Mam malogy, 63: 23–32.

KUNZ, T. H., R. A. ADAMS, and W. R. HOOD. 2009. Methods

for assessing postnatal growth and development of bats.

Pp. 273–324, in Ecological and behavioral methods for the

study of bats, 2nd edition (T. H. KUNZ and S. PARSONS, eds.).

The Johns Hopkins University Press, Baltimore, 901 pp.

KUNZ, T. H., and W. R. HOOD. 2000. Parental care and postna-

tal growth in the Chiroptera. Pp. 415–468, in Reproductive

biology of bats (E. G. CRICHTON and P. H. KRUTZSCH, eds.).

Academic Press, London, 510 pp.

KUNZ, T. H., and S. K. ROBSON. 1995. Postnatal growth and

development in the Mexican free-tailed bat (Tadarida bra -siliensis mexicana): birth size, growth rates, and age estima-

tion. Journal of Mammalogy, 76: 769–783.

LIU, Y., L. R. JIN, W. METZNER and J. FENG. 2009a. Postnatal

Postnatal growth and age estimation in Rhinolophus marshalli 109

growth and age estimation in big-footed myotis, Myotismacrodactylus. Acta Chiropterologica, 11: 105–111.

LIU, Y., T. L. JIANG, S. BERQUIST, and J. FENG. 2009b. Vocal

characters and wing morphology of Rhinolophus marshallifrom Tiantang Cave, Guangxi Province, China. Mammalia,

73: 373–376.

MCLEAN, J. A., and J. R. SPEAKMAN. 2000. Morphological

changes during postnatal growth and reproduction in the

brown long-eared bat Plecotus auritus: implications for

wing loading and predicted flight performance. Journal of

Natural History, 34: 773–791.

RAGHURAM, H., and G. MARIMUTHU. 2007. Maternal feeding of

offspring with vertebrate prey in captive Indian false vam-

pire bat, Megaderma lyra. Acta Chiropterologica, 9: 437–443.

RAJAN, K. E., and G. MARIMUTHU. 1999. Postnatal growth and

age estimation in the Indian false vampire bat (Megadermalyra). Journal of Zoology (London), 248: 529–534.

REITER, G. 2004. Postnatal growth and reproductive biology of

Rhinolophus hipposideros (Chiroptera: Rhinolophidae).

Jour nal of Zoology (London), 262: 231–241.

SHARIFI, M. 2004a. Postnatal growth and age estimation in the

Mehelys horseshoe bat (Rhinolophus mehelyi). Acta Chiro -

pterologica, 6: 155–161.

SHARIFI, M. 2004b. Postnatal growth in Myotis blythii (Chiro -

ptera, Vespertilionidae). Mammalia, 68: 283–289.

SIMMONS, N. B. 2005. Order Chiroptera. Pp. 312–529, inMam mal species of the World: a taxonomic and geograph-

ic reference, 3rd edition (D. E. WILSON and D. M. REEDER,

eds.). The Johns Hopkins University Press, Baltimore,

2142 pp.

STERN, A. A., and T. H. KUNZ. 1998. Intraspecific variation in

postnatal growth in the greater spear-nosed bat. Journal of

Mammalogy, 79: 755–763.

ZHANG, L. B., G. JONES, J. S. ZHANG, G. J. ZHU, S. PARSONS, S.

J. ROSSITER, and S. Y. ZHANG. 2009. Recent surveys of bats

(Mammalia: Chiroptera) from China. I. Rhinolophidae and

Hipposideridae. Acta Chiropterologica, 11: 71–88.

ZULLINGER, E. M., R. E. RICKLEFS, K. H. REDFORD and G. M.

MACE. 1984. Fitting sigmoidal equations to mammalian

growth curves. Journal of Mammalogy, 65: 607–636.

110 L. Jin, L. Bo, K. Sun, Y. Liu, and J. Pan Ho

Received 05 November 2010, accepted 11 April 2011