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Age Estimates for a Population of the Indian Tree Frog Polypedates Maculatus (GRAY, 1833) (Anura: Rhacophoridae)

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Age Estimates for a Population of the Indian Tree Frog Polypedates Maculatus (GRAY, 1833) (Anura: Rhacophoridae) HERPETOZOA 21 (1/2): 31 - 40 31 Wien, 30. Juni 2008 Age estimates for a population of the Indian Tree Frog Polypedates maculatus (GRAY, 1833) (Anura: Rhacophoridae) Altersschätzungen an einer Population des Gefleckten Ruderfrosches Polypedates maculatus (GRAY, 1833) (Anura: Rhacophoridae) PRAVATI K. MAHAPATRA & SUSAMA NAYAK & SUSHIL K. DUTTA KURZFASSUNG Die Arbeit berichtet über die skelettochronologische Abschätzung von Alter und Lebenserwartung sowie des Alters bei Eintritt der Geschlechtsreife bzw. Fortpflanzung bei Polypedates maculatus (GRAY, 1833). Die Auf- sammlung der Frösche (n = 76) erfolgte von Juni bis September in den Jahren 2005 und 2006. Bei jedem Tier wur- den die Kopf-Rumpf-Länge gemessen und die 4. Zehe des linken Hinterbeines unter lokaler Betäubung abgetrennt und in 70 %igem Äthanol fixiert. Acht Tiere aus unterschiedlichen Größenklassen wurden getötet und dienten zur Bestimmung des Reifegrades der Gonaden; an ihnen wurde auch die Histologie der Phalangen- und Langknochen Humerus und Femur untersucht. Nach äußeren und gonadalen Merkmalen wurden drei Gruppen von Fröschen unterschieden - unreife Jungtiere (Gruppe I), geschlechtsreife Männchen und Weibchen (Gruppe II) und Pärchen in Umklammerung (Gruppe III) Die gewonnenen Phalangen- und Langknochen wurden in 10 %iger EDTA ent- kalkt und zur Anfertigung histologischer Schnitte weiterverarbeitet. Ein bis sechs Wachstumsringe, jeder aus einer Wachstumszone und einer Zone verminderten Wachstums (LAG) bestehend, waren bei geschlechtsreifen Fröschen ausgebildet. Da jedes Jahr ein Wachstumsring hinzukommt, ergibt sich bei dieser Froschart in natürlichen Popu- lationen ein Höchstalter von sieben Jahren . Unreife Jungtiere zeigten keine LAGs. Bei Pärchen in Umklam- merung hatten die Individuen eine bis fünf LAGs. Dabei waren die Weibchen immer größer (Kopf-Rumpf-Länge) als ihre Partner, die ihrerseits entweder ebensoalt wie oder jünger als die zugehörigen Weibchen waren. ABSTRACT Skeletochronological estimation of age, longevity, age at sexual maturity and breeding of the Indian Tree Frog Polypedates maculatus (GRAY, 1833) is described in this paper. The frogs (n=76) were collectedduring the months of June – September in the years 2005 – 2006. The snout-vent length of each frog was recorded. The 4th toe of the left hind limb of each frog was clipped under anesthesia and fixed in 70% ethanol. Eight frogs belonging to different size groups were sacrificed to determine the gonadal status. The same frogs were utilized to study the histology of both long (humerus and femur) and phalangeal bones. Based on the external morphology and gonadal status, the frogs were divided into three groups i.e. the immatures (Group-I), mature males and females (Group-II) and amplecting pairs (Group-III). Subsequently, phalanges and long bones were decalcified in 10% EDTA and pro- cessed for histology. One to six growth rings consisting of growth zones and lines of arrested growths (LAGs) were observed in mature frogs. As a single growth ring is added every year, maximum age of this species is considered to be seven years in natural populations. Immature frogs did not show any LAG. The amplecting pairs showed one to five LAGs and the females were always larger in size (snout-vent length) than the corresponding male partners. Among the amplecting pairs, males were either of the same age group or younger than their female partners. KEYWORDS Amphibia: Anura: Rhacophoridae: Polypedates maculatus, lines of arrested growth, age, growth pattern, growth rings, longevity, population ecology, India INTRODUCTION Skeletochronology is considered to be KUMBAR & PANCHARATNA 2001a). In long the most reliable and powerful tool to esti- bones as well as phalanges, amphibians and mate the age and longevity of amphibians reptiles develop growth marks in which (CASTANET & SMIRINA 1990; SMIRINA 1994; broader zones correspond to periods of fast 32 P. K. MAHAPATRA & S. NAYAK & S. K. DUTTA Table 1: Snout-vent length (SVL, mm) and number of LAGs in different groups of Polypedates maculatus (GRAY, 1833). n – number of frogs; SD – Standard deviation. Tab. 1: Kopf-Rumpf-Länge (SVL, mm) und Anzahl der Linien verminderten Wachstums (LAGs) in ver- schiedenen Altersgruppen von Polypedates maculatus (GRAY, 1833). n – Stichprobengröße; SD – Standardabwei- chung; Female – Weibchen; Male – Männchen. Age group SVL (mm) Number of LAGs / Anzahl LAGs n Mean ± SD (Range) Median / Mode (Range) Altersgruppe Mittelwert ± SD (Spannweite) Median / Modalwert (Spannweite) Immature / Jungtiere 16 33.03 ± 9.19 (15.0 – 50.0) 0 / 0 (0 – 0) Mature / Geschlechtsreife Female 26 61.32 ± 10.67 (41.0 – 83.0) 3 / 1 (1 – 6) Male 24 44.61 ± 3.79 (35.0 – 53.0) 2 / 2 (1 – 4) Amplecting Pairs / Verpaarte Female 5 59.9 ± 3.87 (56 – 65.5) 4 / 5 (1 – 5) Male 5 45.1 ± 2.45 (42 – 47.5) 3 / 3 (1 – 3) growth and lines of arrested growth (LAGs) CHARATNA & DESPANDE 2003). Skeleto- to resting periods (CASTANET & SMIRINA chronological estimation of age of only two 1990; SMIRINA 1994; KUMBAR & PANCHAR- anurans inhabiting eastern India has been ATNA 2001a). So, one broad zone (growth reported (NAYAK et al. 2007, 2008). So, in ring) plus its enveloping LAG represents the present study an attempt has been made one year of growth, which corresponds to to investigate the age, longevity, age at sex- the age of the individual (HALLIDAY & ual maturity and breeding in a natural popu- VERRELL 1988; CASTANET & SMIRINA 1990; lation of the Indian rhacophorid tree frog, SMIRINA 1994). There are few reports on Polypedates maculatus (GRAY, 1833) inhab- skeletochronological estimation of age in iting Bhubaneswar (20°18’ N, 85°50’E), some south Indian anurans (KUMBAR & eastern India. PANCHARATNA 2001a, 2001b, 2002; PAN- MATERIALS AND METHODS During the course of study (June 2005 and amplecting pairs (Group III, n = 5 + 5), to September 2006), 76 specimens of respectively. Both long bones and toes were Polypedates maculatus (GRAY, 1833) meas- water rinsed for one hour and decalcified in uring 15.0 to 83.0 mm in snout-vent length 10% EDTA (ethylene diamine tetra-acetic (SVL) were collected from various locali- acid) solution for 24-48 h, depending on the ties of Bhubaneswar city, Orissa, eastern size of the bone. After decalcification, the India. Following capture, the frogs were bones were rinsed thoroughly for 24 h in anaesthetized with ether and the SVL was running tap water. Following rinsing, the recorded. The 4th toe of the left hind limb bones were dehydrated and processed for was cut off from each frog and fixed in 70% paraffin block preparation. Serial sections ethanol. To study the gonadal status, eight ranging from 8 to 10 µm were cut using a frogs of different size groups were sacri- rotary microtome. The sections were ficed. The same specimens were fixed in stained in Delafield’s haematoxylin and eo- 70% ethanol for study of histology of both sin and examined under a compound micro- long (femur and humerus) and phalangeal scope. Photographs were obtained from bones. The remaining 68 frogs were appropriate sections using an Olympus® released to the locality of their capture. OIC 04194 microscope and Pentax® K1000 Based on the size, secondary sex characters camera. The relationship between body size and gonadal status, the specimens were (SVL) and age was assessed by drawing divided into three groups, i.e. immature scattered plots and calculating correlation males and females (Group I, n = 16), mature coefficients (Pearson’s r’) (STEEL & TORRIE males and females (Group II, n = 26 + 24) 1980). Age estimates in Polypedates maculatus (GRAY, 1833) 33 Table 2: Snout-vent length (SVL, mm) and number of LAGs in 10 individuals of five amplecting pairs of Polypedates maculatus (GRAY, 1833). F – Female; M – Male. Tab. 2: Kopf-Rumpf-Länge (SVL, mm) und Anzahl der Linien verminderten Wachstums (LAGs) bei fünf Pärchen (10 Individuen) von Polypedates maculatus (GRAY, 1833). F – Weibchen; M – Männchen. Amplecting pair # SVL (mm) Number of LAGs / Anzahl LAGs Pärchen Nr. F / M F / M 1 62.0 / 43.0 1 / 1 2 56.0 / 47.0 2 / 2 3 59.0 / 46.0 5 / 3 4 57.0 / 42.0 5 / 3 5 65.5 / 47.5 4 / 3 Growth pattern in body size (SVL) was ‘b’ is the growth rate, ‘c’ is the constant and analyzed by fitting the logistic model ‘t’ is the time (age) in years. We used the L = a / (1 + bect), where ‘L’ is the average software package SPSS® to compute and SVL at a given age t, ‘a’ is the asymptotic visualize the logistic regression function. SVL; ‘e’ is the base of natural logarithms, RESULTS Cumulative data on size (SVL) and of femur of the smallest frog (SVL 15.0 number of LAGs present in different groups mm) showed a thin cortex (Fig. 1A). The is given in Table 1. The matured males were endosteal and periosteal layers were indis- comparatively smaller in size than the fe- tinct. The marrow cavity was filled with males in the natural population. Similarly, vacuolated cells and there was no LAG in among the amplecting pairs the SVLs of the matrix. The cross section of the femur males were found to be less than in the cor- bone of another immature frog (SVL 42.0 responding amplecting females (Table 2). mm) showed a comparatively broader cor- The histological structure of both long tex than the smallest captured frog, with bones and the phalanges showed a central evenly scattered osteocytes (Fig. 1B) and marrow cavity (mc) lined by an endosteal showed no LAG in the cortex. layer (e) (Figs. 1A, 1B). The middle layer In group II (matured) frogs, one to (cortex, c) consisted of a matrix and evenly four LAGs were observed in males (Table 1 scattered osteocytes (o). The cortex was and Figs. 1C, 2B, 2C), whereas one to six covered by an outer periosteal layer (p). In LAGs were found in females (Figs. 1D and the mature frogs, a series of concentric 2A, 2D).
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