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Postnatal Growth and Age Estimation in the Ashy Leaf-Nosed Bat, HipposideroscineraceusAuthor(s): Longru Jin, Aiqing Lin, Keping Sun, Ying Liu and Jiang FengSource: Acta Chiropterologica, 12(1):155-160. 2010.Published By: Museum and Institute of Zoology, Polish Academy of SciencesDOI: http://dx.doi.org/10.3161/150811010X504653URL: http://www.bioone.org/doi/full/10.3161/150811010X504653

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INTRODUCTION

The time during which a young mammal devel-

ops appropriate sensory and locomotor skills neces-

sary to become independent from its mother is de-

fined as the postnatal growth period (Baptista et al.,2000). Many researchers have studied this period

with respect to changes in behavior, physiology and

ecology to investigate life history traits and to esti-

mate maternal investment (Kunz and Hood, 2000).

There have been numerous studies on postnatal de-

velopment of bats under both natural (Hoying and

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

verri and Kunz, 2006; Liu et al., 2009; Wei et al.,2009) 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 measurements of body mass,

forearm length, the total length of the epiphyseal

gap of the fourth metacarpal-phalangeal joint (De

Fanis and Jones, 1995; Hoying and Kunz, 1998; Liu

et al., 2009), and changes in tooth development

(Anthony, 1988) can all be used to estimate the age

of bats during the early postnatal period. Forearm

length is the most accurately measured and reliable

character for estimating age during the early linear

growth period of bats. However, 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 nutri-

tional intake and energy expenditure over the short

term, and to daily water flux (Kunz, 1987; 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 devel-

opment vary among species and families of bats.

Acta Chiropterologica, 12(1): 155–160, 2010PL ISSN 1508-1109 © Museum and Institute of Zoology PAS

doi: 10.3161/150811010X504653

Postnatal growth and age estimation in the ashy leaf-nosed bat,

Hipposideros cineraceus

LONGRU JIN1, 2, AIQING LIN1, 2, KEPING SUN1, 2, YING LIU1, 2, and JIANG FENG1, 2, 3

1Key Laboratory for Wetland Ecology and Vegetation Restoration of National Environmental Protection, Northeast Normal University, Changchun 130024, China

2Key Laboratory of Vegetation Ecology of Education Ministry, Institute of Grassland Science, Northeast Normal University,Changchun 130024, China

3Corresponding author: E-mail: fengj@nenu.edu.cn

We quantified changes in body mass, forearm length, and the total length of the epiphyseal gap of the fourth metacarpal-phalangeal

joint of the ashy leaf-nosed bat (Hipposideros cineraceus) based on mark-recapture data obtained in Fangkong Cave in Hekou,

Yunnan Province, China. We used these data to develop empirical growth curves, to derive growth rates, to establish age-predictive

equations, and to compare growth parameters based on three nonlinear growth models. Forearm length and body mass of pups

followed a linear pattern of growth until day 17, with mean growth rates of 0.81 mm/day and 0.09 g/day, respectively and thereafter

their growth rates increased more slowly. The length of the epiphyseal gap initially increased linearly up to day 13 and then

decreased linearly at a mean rate of 0.07 mm/day until day 37. An equation for estimating age based on forearm length was valid

when this dimension was ≤ 27.6 mm, whereas the equation based on the length of the epiphyseal gap was valid for forearm lengths

≥ 24.3 mm. Together, these two equations permit estimation of the age of H. cineraceus pups between 1 and 37 days. 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: Hipposideros cineraceus, postnatal growth, age estimation, nonlinear growth models

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 independent

of body size and duration of the postnatal growth pe-

riod (Kunz and Robson, 1995). However, compara-

tive studies among different taxa should be based on

the same models (Zullinger et al., 1984).

In this paper, we investigated the postnatal de -

velopment of the ashy leaf-nosed bat, Hipposi derosc ineraceus. This is a widespread species of bat,

rang ing through Pakistan India, Burma, Thailand,

Laos, Vietnam, Sumatra and Borneo (Simmons,

2005). Recently, it has also been found in China

(Zhang et al., 2009). However, little information is

available on its external characters at birth and

during postnatal growth (Gould, 1979). Thus, the

aim of this paper is to provide such background data

on the pattern of postnatal growth for a free-rang -

ing population of H. cineraceus and to derive age-

specific growth equations based on a quantitative

analysis of forearm length and the epiphyseal gap.

MATERIALS AND METHODS

The study was conducted between May and July 2009 in

Fangkong Cave (22°36’N, 103°51’E; ca. 100 m long, 2 m wide,

and 2.2 m high) in Hekou County, Yunnan Province, China. This

cave housed a mixed colony of about 200 H. cineraceus,

150 Hipposideros pomona, and 50 Hipposideros larvatus. The

surrounding area was covered by dense forest of mostly Heveabrasiliensis.

Prior investigation indicated that the females of H. cinera -ceus in this area give birth from late April to early May. There -

fore, we checked the cave daily from about 15 days before par-

turition. We hand-captured neonates immediately following the

nightly emergence of adults. Neonates with an attached umbili-

cal cord were assumed to be one day old (Kunz and Robson,

1995). Following determination of the sex of pups, a uniquely

number ed plastic split-ring band (Porzana Ltd., UK) was

placed on the forearm of each bat for individual identification.

In total, 48 neonates with umbilical cords were marked at the

first day.

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

tronic balance (ProScale LC-50, America). Forearm length was

measured to the nearest 0.01 mm with a digital vernier caliper

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

seal gap was measured to the nearest 0.01 mm using the caliper

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

plastic sheet illuminated from below with a strong torch to

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

vernier calipers, individual measurements were repeated five

times and the means used in the analysis.

To limit disturbance and possible abandonment, marking of

young and measuring morphological characters of pups took

place while adults were out foraging and the whole task were

completed within 1.5 h. As soon as all young had been measured

and weighed, they were returned to the sites, as near to the orig-

inal roost as possible, before any females returned. The cave

was visited three times in the first week and thereafter every

four days (Table 1). Visits were made until the pups were

observed to begin foraging trips, by which time it became

difficult to capture them even using hand nets and mist nets. The

capture of bats was performed with the permission of the local

government.

Linear regression equations were derived to predict age

from pooled data for forearm length (1–17 days) and total epi-

physeal gap (13–37 days). To derive age-predictive equations

from these measurements, the axes on the growth curves were

reversed and age was treated as the dependent variable (Kunz

and Anthony, 1982). Ninety-five percent confidence intervals

and prediction limits were plotted for the regression equa-

tions for forearm length and total gap. In addition, growth data

for young bats were fitted to the following three models: the

logistic equation, the Gompertz equation, and the von Berta -

lanffy equation (Zullinger et al., 1984). The equations were as

follows:

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

2. Gompertz: W = A exp (- exp (- K (t - I)))3. 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 neo -

nate 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 different models were compared by the goodness of fit ob-

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

analyses were conducted using SPSS 15.0.

RESULTS

The first newborn H. cineraceus with attached

umbilical cord was found on May 8th and the last

one on May 24th 2009. Each female observed deliv-

ered single offspring. At birth, the young were naked

and pink with closed eyes, folded ears, and decidu-

ous teeth. The thumbs and feet were large enough

to enable the young to cling to and to grasp the

mother’s fur at the time of birth. Neonates were born

156 L. Jin, A. Lin, K. Sun, Y. Liu, and J. Feng

TABLE 1. Number of individuals captured on day one and subsequently re-captured on other sampling occasions

ParameterDay of sampling

1 3 5 9 13 17 21 25 29 33 37

Number of re-captured bats 48 38 36 38 37 33 38 30 25 20 14

Marked bats re-captured (%) – 79 75 79 77 69 79 62 52 42 29

with the forearm length averaging 42.2% of the

length in adult females, and 36.1% of the body mass

of adult females. Five days after birth, the short,

fine, soft hair of the pups was distinguishable. Thir -

teen days after birth, most pups’ eyes were com-

pletely open and dark fur similar to that of adults

developed. At this time, their ears were erect. After

25 days, some young were able to flutter and glide

when they were released by hand. After 33 days,

most young bats were observed to fly freely with

gentle turns in the cave. At that time, mean body

mass and forearm length were 81.4% and 95%, of

the corresponding values for adult female. Finally,

37 days after birth, most young bats left the cave for

their first foraging trips.

During the first 17 days body mass (Fig. 1A) and

forearm length (Fig. 1B) increased linearly with

growth rates of 0.09 g/day and 0.81 mm/day, respec-

tively (empirical growth curves from 48 young).

Sub sequently, the growth rates of these two

characters decreased to a relatively constant value.

In contrast, the length of the epiphyseal gap in-

creased linearly until day 13 and then decreased

linearly at a mean rate of 0.07 mm/day (Fig. 1C).

A linear regression equation was derived for

estimating the age of the young bats on the basis of

forearm lengths between 14.6 mm (mean on the 1st

day) and 27.6 mm (up to 17 days). A second

equation estimated the age from the total length

of the epiphyseal gap until day 37. Together,

these two equations allow to predict the age of

H. cineraceus from 1 to 37 days after birth (Figs. 2

and 3).

Based on the coefficient of determination obtain -

ed from the fitted curves, the logistic model appears

to provide the best fit for body mass and forearm

length (Table 2). Body mass and forearm length

indicated asymptotic maximum values of 3.53 g

with a growth rate of 0.12 g/day and 34.69 mm with

a growth rate of 0.10 mm/day, respectively (Table 2).

DISCUSSION

The selection of appropriate methods of data col-

lection, and for the analysis of postnatal growth, is

extremely important because each technique can

significantly influence the estimates of growth pa-

rameters (Kunz and Robson, 1995). The two most

common methods used to assess postnatal growth

in the field are based on mark-recapture (longitudi-

nal) sampling and grab (cross-sectional) sampling.

Bap tista et al. (2000) compared these two sampling

methods for prediction of the age of free-ranging

Postnatal development in Hipposideros cineraceus 157

FIG. 1. Empirical growth curves for A — body mass (n = 357),

B — forearm length (n = 357), and C — total length of the

gap of the fourth metacarpal-phalangeal joint (n = 357) of

H. cineraceus from day 1 to 37. Some points represent more

than one observation

bats. They concluded that the longitudinal method

was the more reliable technique for age estimation.

Under captive conditions, bats are commonly hand-

raised to study their postnatal development in more

detail. However, a marked difference between cap-

tive and free-ranging bats has been reported (Gould,

1971; Buchler, 1980). Captive bats are often fed

an unnatural diet (e.g., mealworms) and may not be

provided with appropriate roosting environments or

adequate space for exercise. Thus the flight costs of

Age (days)

Fore

arm

length

(m

m)

0 10 20 30 40

40

30

20

10

Body m

ass (

g)

4

3

2

1

Length

of

epip

hyseal gap (

mm

)

4

3

2

1

A

B

C

158 L. Jin, A. Lin, K. Sun, Y. Liu, and J. Feng

FIG. 2. Regression line estimating the age of H. cineraceus from

values of forearm length up to 17 days. Some points represent

more than one observation. The predictive equation is valid for

forearm lengths ranging from 14.6 to 27.6 mm. Narrow and

wide bands indicate 95% confidence and prediction intervals,

respectively; age (days) = 1.22 × length of forearm - 16.76

(r2 = 0.99, n = 230, P < 0.01)

FIG. 3. Regression line estimating the age of H. cineraceus from

the values of the total length of the gap of the fourth metacarpal-

phalangeal joint at 13–37 days. Some points represent more

than one observation. The predictive equation is valid for

forearm lengths ranging from 24.3 to 33.3 mm. Narrow and

wide bands indicate 95% confidence and prediction intervals,

respectively; age (days) = 55.8 - (12.97 × length of total gap)

(r2 = 0.91, n = 197, P < 0.01)

than reported values of ca. 25% (20–30%) of adult

female mass for most pups of insectivorous bats at

birth (Kurta and Kunz, 1987). The explanation

appears to be that litter mass among bats is highly

correlated with maternal body mass in an allometric

relationship, with smaller bats having relatively

large offspring and large bats having relatively small

offspring (Hayssen and Kunz, 1996). Large size

of bats at birth is generally associated with an

advanced stage of development and is believed to

aid the conservation of heat generated to maintain

constant body temperature (Reiter, 2004). This sup -

ports an advanced state of neuromuscular devel op -

ment at birth (Kurta and Kunz, 1987).

Sustained flight of young bats occurred in 33

day-old pups, by which time they had achieved

81.4% of adult body mass and 95% of adult forearm

proportions. This is similar to juveniles of other in-

sectivorous 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 small body size and large

skeletal size at the onset of flight lowers wing load-

ing and this in turn increases maneuverability, and

decreases the cost of flight, at a time when the young

bats are learning how to detect and capture flying in-

sects (Hughes et al., 1995).

The pattern of the postnatal growth in H. ciner-aceus is consistent with that of many species of bats

(Hoying and Kunz, 1998; Stern and Kunz, 1998;

Rajan and Marimuthu, 1999; Hood et al., 2002;

adults are also likely to be lower than in the wild. In

fact, recent studies have indicated no significant dif-

ference in the growth patterns of young held in cap-

tivity compared with young growing under natural

conditions (e.g., Rousettus leschenaultii — Elan go -

van et al., 2002; Megaderma lyra — Rajan and Ma -

ri muthu, 1999). However, these studies were re-

stricted to fruit and vampire bats. The ashy leaf-

nosed bats used in this study are insectivorous and,

as their pups are easily recaptured following the

nightly emergence of the adult bats, mark-recapture

sampling was used to obtain data for study of their

postnatal development.

The characteristics of neonates of the ashy leaf-

nosed bat, including closed eyes and naked skin, are

similar to most other bat species. However, the eyes

of H. cineraceus did not completely open until 13

days, which is slower than in most other insectivo-

rous bats, e.g., ca. five days in Myotis macrodactylus(Liu et al., 2009) and Rhinolophus mehelyi (Sharifi,

2004a). However, the eyes of Hipposideros terasen-sis (Chen et al., 2002) and Hipposideros speoris(Habersetzer and Marimuthu, 1986) pups only

opened after two weeks. In the field, we also found

that the eyes of H. pomona and H. larvatus pups

opened after 10 days. Thus, it is possible that this

phenomenon is common to the Hipposideridae.

The mean mass at birth of neonates of H. cinera -ceus was 36.1 % of the mean mass of female adults

and their mean forearm length was 42.2% of the

maternal length. These values are somewhat higher

Length of epiphyseal gap (mm)

Age (

days)

1 2 3 4

40

30

20

10

Forearm length (mm)

Age (

days)

12 16 20 24 28

20

15

10

5

0

Reiter, 2004). Forearm length and body mass in-

creased linearly during the early preflight period and

the total length of the gap of the fourth metacarpal-

phalangeal joint decreased linearly during the late

period. According to age estimations of H. cinera -ceus, forearm length is useful in determining the age

of the young during the preflight stage when its

growth is linear. The total gap length of the fourth

metacarpal-phalangeal joint is useful as an addition-

al parameter for age estimation during the postflight

stage, when growth of the forearm becomes non-lin-

ear. By using measurements of the total epiphyseal

gap and forearm length, it is possible to reliably

estimate the age of young H. cineraceus from 1 to

37 days. Linear equations can be used effectively to

describe rates of change during different phases of

the postnatal period and they also can be valuable

for assigning ages to bats during the postnatal period

(Anthony, 1988; Bohn et al., 2007). However, post -

natal growth rates are influenced by climate, food

supply, habitat, latitude, maternal factors, and social

environment (Cumming and Bernard, 1997; Hoy-

ing and Kunz, 1998; Hood et al., 2002; Dietz et al.,2007). Thus, we encourage researchers to compare

the changes in different years and to generate geo-

graphically specific equations whenever possible.

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.

It is computationally simple and has biological rele-

vance (Kurta and Kunz 1987; Kunz and Robson,

1995; Krochmal and Sparks, 2007). Therefore, as

in other bat species, e.g., Pipistrellus pipistrellus(Boyd and Myhill 1987), Plecotus auritus (De Fanis

and Jones, 1995), Tadarida brasiliensis (Kunz and

Rob son, 1995), Myotis septentrionalis and Myotisluci fugus (Krochmal and Sparks, 2007), the logistic

model provided the best fit to our empirical data for

H. cineraceus.

ACKNOWLEDGEMENTS

This study was financed by the National Natural Science

Foundation of China (Grant No. 30770361, 30870371), the

National Grand Fundamental Research 973 Program of China

(2009CB426305) and the Doctoral Foundation of Ministry of

Education (20060200007). We thank Zheng Liu for his invalu-

able field assistance.

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Postnatal development in Hipposideros cineraceus 159

TABLE 2. Growth parameters in H. cineraceus derived from the logistic, Gompertz, and von Bertalanffy growth models. Parameters:

A — asymptotic value of forearm length (cm) or body mass (g), K — growth rate constant, I — inflection point, SE — standard

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Body mass versus age Forearm length versus age

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Model sum Estimate SE CV (%)

Model sum

of squares of squares

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K 0.12 0.003 2.52 0.10 0.001 0.98

I 3.42 0.145 4.24 4.32 0.077 1.78

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K 0.09 0.003 3.37 0.07 0.001 1.35

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Received 30 September 2009, accepted 31 March 2010