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Analysis of growth in apopulation of Liolaemusoccipitalis Boul. 1885, fromthe Coastal Sand‐dunes ofTramandai, RS, Brazil (Reptilia,Tropiduridae)Laura Verrastro a & Ligia Krause aa Departamento de Zoologia, Instituto deBiociencias Universidade Federal do Rio Grande doSul, Av. Paulo Gama s/n, 90049, Porto Alegre, RS,BrazilPublished online: 19 Nov 2008.

To cite this article: Laura Verrastro & Ligia Krause (1994) Analysis of growth ina population of Liolaemus occipitalis Boul. 1885, from the Coastal Sand‐dunes ofTramandai, RS, Brazil (Reptilia, Tropiduridae), Studies on Neotropical Fauna andEnvironment, 29:2, 99-111, DOI: 10.1080/01650529409360922

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Studies on Neotropical Fauna and Environment 0165-0521/94/2902-0099$6.00Vol. 29 (1994), No. 2, pp. 99-111 © Swets & Zeitlinger

Analysis of Growth in a Population of Liolaemusoccipitalis Boul. 1885, from the Coastal Sand-dunesof Tramandai, RS, Brazil (Reptilia, Tropiduridae)

Laura VERRASTRO and Ligia KRAUSE

VERRASTRO, Laura and Ligia KRAUSE (1994): Analysis of Growth in aPopulation of Liolaemus occipitalis Boul. 1885, from the Coastal Sand-dunes of Tramandai, RS Brazil (Reptilia, Tropiduridae).Studies on Neotropical Fauna and Environment 29, pp. 99-111.

The growth of the species Liolaemus occipitalis was analysed in a coastalpopulation of Rio Grande do Sul. The comparative analysis of growth ratewas carried out between 32 males and 45 females throughout one year.Taking into consideration the well-defined seasonality of this temperatezone, the ontogenetic development (size and weight) was observed. Sexualdimorphism was evident from the viewpoint of size, females always beingsmaller. The annual growth curve of the individuals from this population issimilar to that of lizards from temperate zones, with maximum rates inspring and growth rates close to zero in winter. Among the species of Liolaemusstudied so far, L. occipitalis proves to be one of the smallest.

Laura Verrastro, Ligia Krause: Departamento de Zoologia, Instituto deBiociencias Universidade Federal do Rio Grande do Sul, Av. Paulo Gama s/n, 90049 Porto Alegre, RS, Brazil.

The genus Liolaemus Wiegmann, 1834, is quite diversified, with approximately80 species (Donoso-Barros, 1966) distributed between the 25th and the 55thparallels in South America. In Brazil it is represented by the species L. lutzae andL. occipitalis. The former is limited to the sandbank of Rio de Janeiro (Rocha,1985b) and the latter to the coast of Rio Grande do Sul, with two citations for thestate of Santa Catarina (Müller and Steineger, 1977).

Studies carried out on L. occipitalis are almost exclusively of a taxonomical,biogeographical and osteologic character. Among them we can cite works byVanzolini and Ab'Saber (1968), Müller and Steineger (1977), Gudynas (1981),Keller and Krause (1988), Simöes-Lopes and Krause (1986).

Given the abundance of this lizard in our beaches and the lack of informationabout its biology, it was the purpose of our work to study its growth. To do so, wefocused mainly on determining snout-vent length (SVL) and weight, analysingthe growth rate both in relation to ontogenetic development and in relation to thefour seasons of the year.

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100 LAURA VERRASTRO AND LIGIA KRAUSE

Description of the Site

The study took place in Jardim Atlantico Beach (30°40 'S and 10° 50 'W), in the municipality ofTramandai. The site is 1000 m far from the ocean and approximately 500 m east from the GentilLagoon.

The area is made up of mobile dunes on Holocene soil and is included in the coastal sandy region(Delaney, 1965 and Willwock, 1973) which geomorphologically belongs to the outer coastal plains(IBGE, 1986).

Within the chosen area there is a central depression intercalated by small elevations of sandylittle mounds 20 cm to 50 cm high. When a rainy period occurs, temporary floodings are observeddue to the elevation of the phreatic water-table.

Vegetation is sparse, with a covering of no more than 5%. It is formed exclusively by psammophileherbaceous plants, usually of the family Gramineae. Panicum racemosum is the most abundantspecies, forming aggregates almost homogeneous in large extensions, thriving especially on top ofthe little mounds.

Methodology

The area of study was established when a rectangle was delineated from the line of posts alreadyexistent there and numbered wooden stakes of 2.5 cm x 2.5 cm x 50 cm were distributed to form aright angle with that first line of posts. Two rows of stakes were then placed on the right side and onerow on the left side, with a 30-meter distance between them. Thus an area of 400 m x 150 m wasdetermined (Fig. 1).

To obtain the biométrie data it was used the method of capture, marking, and recapture. Thesedata were collected for one year, between March 1987 and March 1988. During the first semester, atime of lower temperatures (until Aug. 1987), visits to the site were monthly. During the secondsemester (from Sept. 1987 on), visits took place every two weeks, taking into consideration thislizard's higher activity in the warmer months.

The individuals collected were taken to the laboratory to be marked and for us to obtain thebiométrie data: weight (in grams) and length of body (snout-vent length = SVL) and tail (both inmillimeters). Marking was accomplished with the amputation of the last unguinal phalanx followedby a sequence of numbering.

Growth rate was estimated by means of applying the formula used by Van Devender (1978). Theestimate included those individuals that had been captured at least twice at intervals of 30 to 100days, during spring and summer. This is because Van Devender believes data obtained at intervals ofless than 30 days could produce deviations, for the increase in growth could be relatively smaller thanthe mean error; at intervals of more than 100 days, it could reflect seasonality.

Results and Discussion

In the analysis of growth some items were examined which were considered tobe relevant for a better understanding of the development of this species, whosebiology was unknown up to now.

At first, the size, length and weight were determined for this population andthe information provided by these data was analysed.

Secondly, a detailed analysis of the growth rate was carried out, basicallyaccording to two approaches: (a) in relation to ontogenetic development and (b)in relation to the seasons of the year.

Table 1 shows the average, minimum, and maximum SVLs found in adults ofthis population during the period of study. For this analysis the minimum size ofan adult was considered to be the size of the individual which achieved sexualmaturation, according to Verrastro (1991). Average SVL was estimated using all

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GROWTH IN A POPULATION OF LIOLAEMUS OCCIPITALIS 101

Dun»1

Fig. 1. Establishing and marking the main site, where capture, marking and recapture of the lizardswere done. P = posts already existent in the site; E = numbered stakes put there for this study;Dunes = natural delimitation.

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102 LAURA VERRASTRO AND LIGIA KRAUSE

Table I. Differentiated sizes of males and females (mm).Minimum size of first adult stage determined by sexual maturation (Min.s.)- Average size ofan adult estimated through the mean value (Aver.s.). Maximum size represented by thelargest individual found in the population (Max.s.). Average weight of an adult estimatedthrough the mean value (Aver.w.).N = number of individuals

Min.s. Aver.s. Max.s. Aver.w. N

? 45 53.2 63.7 4.9 115

6 50 60.2 71.8 7.2 76

adult individuals found in the population under study, and the results were 53.2mm for females and 60.2 mm for males. Maximum and minimum SVLs weregiven by the largest and the smallest individuals found, respectively. Among thefemales, the highest value found was 63.7 mm and among the males it was 71.8mm. The smallest individual of the population measured 27.7 mm (a hatchling).

When compared to other species of Liolaemus, L. occipitalis presents thelowest values of size ever observed for this genus up to now. Rocha (1985a)finds out an average size of 65-75 mm for L. lutzae and Fuentes (1977) deter-mines a SVL of 84.3 mm for males and 75.7 mm for females of the species L.nigromaculatus.

Following the same methodology employed for SVL, average weight wasestablished (see Table 1); results were 4.9 g for females and 7.2 g for males. Thelowest weight recorded among the hatchlings was 0.5 g. Given the many vari-ables which affect the development of the individuals before they reach adult-hood, maximum weight was disregarded, for it is believed that it does not con-tribute significant data.

As it can be seen, males reach average and maximum sizes which are alwayssignificantly greater than those of the females, thus reflecting a clear sexualdimorphism. This type of dimorphism can be observed for most lizards.

In the case of L. occipitalis, females reach adulthood at 45 mm and males at50 mm, within the same period of time for both sexes. This occurs because thespeed of growth for each sex is different, generally faster in males, as it hasalready been mentioned by Andrews (1982), among others. Andrews also statesthat in most cases when males are bigger than females, the juvenile males growmore rapidly than the juvenile females.

When the relation between weight and SVL is analysed for L. occipitalis, as itwas done for other species, a high correlation becomes evident both for males(r = 0.95 p < 0.001) (Fig. 2) and females (r = 0.88 p < 0.001) (Fig. 3). This corre-lation, slightly lower for the females, is probably caused by the presence of eggsin the oviducts during the period of reproduction. Results show that weightincreased practically in the same proportion as length.

However, the relation found in this study is not a constant for the specimensof this family. For instance, Davis (1968), studying Sceloporus occidentalis,found a relation between these parameters where juveniles increase their SVL

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GROWTH IN A POPULATION OF LIOLAEMUS OCCIP1TALIS 103

3=

12

11

10

9

S

7

e

s

4

2

2

n •

y=-11.20 + 0.30X

SO S4 70

Fig. 2. Relation between SVL (mm) and weight (g) of males of L. occipitalis.

twice as much as they increase their weight, whereas Gennaro (1974), studyingHolbrookia m. maculata, though finding a higher value for length increase thanfor weight increase, the first parameter is not double the second, as in Davis'sexample. This makes us think that the difference in the proportionality betweengrowth and weight of Liolaemus occipitalis, in relation to other species, appar-ently reveals that this species would invest less in body growth during its juve-nile stage than other species of Tropiduridae. In those species of short life span,as it seems to be the case here (Verrastro, 1991), it is expected that juvenileswould reach adult size as quickly as possible, thus reducing the risks of prédationand becoming reproductively active sooner. It would be interesting to analyse, inthe future, whether this result is due to a characteristic of the species or to factorsinherent to this population.

On the other hand, on analysing the growth rate, which reflects the changesthat took place with time, a lengthy discussion is made possible on the type ofdevelopment this species has and its relation to ecology and biology when com-pared to other lizards.

Therefore, considering its ontogenetic development, a negative relation isclearly observed between growth rate and the individuals' SVL (Figs. 4-5). Thiscorroborates the idea that young individuals present higher growth rates thanadults.

This relation is even more evident for males (r = - 0.85 p < 0.001) than for

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104 LAURA VERRASTRO AND LIGIA KRAUSE

45 47 43 51 S3 SS S7 S9 61 63

Fig. 3. Relation between SVL (mm) and weight (g) of females of L. occipitalis.

females (r = - 0.64 p < 0.001), corroborating what was previously discussedwhen analyzing the relation SVL vs. weight. Again, in relation to the females,the correlation is reduced because, spawned in the reproductive period, theydeviate part of their energy to this function and not to growth. Such facts havealso been presented by Fitch (1958) for Cnemidophorus sexlineatus, by Tinkle(1967) for Uta stansburiana, and by Van Devender (1978) in his study of Basiliscusbasiliscus.

Andrews (1982) mentions two reasons which could explain the rapid growthof lizards before they reach adult size: (1) as reproduction in reptiles is seasonal,a slow juvenile growth would delay reproduction; (2) the smaller individuals aresubject to prédation; therefore, a rapid growth of the juveniles would increase thechances of survival.

As Tinkle (1967) wrote, there is a great variation in growth from one indi-vidual to another. However, Andrews (1982) mentions that, generally speaking,reptiles never stop growing; after reaching an asymptotic size, its growth isinsignificant. Andrews states that one of the features of reptilian growth is itsdiscontinuity due to energetic deviations during critical periods: birth, sexualmaturation, and reproduction, or else because of environmental factors. Thus,there are three limitations on growth: (1) size of individual on hatching, (2) sizeat sexual maturation, and (3) maximum body size. According to the observationscarried out during this study of L. occipitalis, it can be stated here that most

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GROWTH IN A POPULATION OF LIOLAEMUS OCCIPITALIS 105

Ioa.o

36 40 44 48 52 56 60 64 68

Fig. 4. Relation between growth rate and SVL (mm) of males of L. occipitalis.

individuals reach a maximum growth rate before they reach sexual maturation.From then on, these rates are reduced until they reach values around 1 mm per

month and even close to zero in larger individuals.An analysis was carried out of some seasonal variations of growth, using for

this purpose the hatchlings that emerged during the one-year period of this study,in order to establish a relation between growth and the seasons of the year.

It was observed, during autumn, that the individuals that hatched in mid-summer (January) had already reached a SVL of approximately 70% of theaverage size they would present when adults (42.2 mm S and 37.4 mm $ ). In thecase of those hatched at the end of the reproductive period (March), the valuesare invariably less than 40 mm (Figs. 6-7).

The curve signals a growth which is gradually slower from autumn to winter.Although data were not obtained in large quantity during winter, the fact iscorroborated by the data of recapture in spring, when the specimens presentedpractically the same size they had in autumn.

Fitch (1958), studying a population of Cnemidophorus sexlineaius, observed,during autumn, individuals that had hatched from different clutches throughoutthe reproductive season. Those from the first clutches reached half the size theywould have in the adult stage before hibernation. Those from late clutches weresmaller when the hibernation period started.

As it was analysed by Andrews (1982), in temperate zones, reptiles present an

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106 LAURA VERRASTRO AND LIGIA KRAUSE

Oa.o

y_ 14.17 - 0.20 x

34 36 28 40 42 44 46 48 SO 52 S4 56

SM. (mm)

Fig. 5. Relation between growth rate and SVL (mm) of females of L. occipitalis.

annual pace of growth, determined by factors both inherent and external. Sea-sonal growth rates are likely to occur, maximum in summer and minimum inwinter.

Moreover, it was observed that juvenile individuals present low growth ratesduring autumn and winter, this showing not a halt in growth but rather a consid-erable decrease of the rate.

This has also been observed by Tinkle (1967) in a study of Uta stansburiana,when it was seen that there is no significant difference between values for winterand values for early spring. Tinkle explains that this would be impossible in casethe individuals depended only on their reserve fat to keep alive. When the cli-mate is favorable, this species feeds and continues growing, though at minimumrates.

In this population of L. occipitalis no characteristic hibernation occurred,although its activity was considerably reduced during winter. Whenever the sunheated the substratum, individuals with some activity could be observed.

On the other hand, starting in spring, a pronounced increase in growth forboth sexes is detected, reaching its maximum in mid-season and early summer.At this time, most of the individuals hatched during the year reach adult size,becoming apt to reproduce. In summer, a decrease is observed in the pace ofgrowth in relation to spring, a fact clearly reflected on the curves of Figures 6and 7.

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GROWTH IN A POPULATION OF LIOLAEMUS OCCIPITALIS 107

75

70

ts

•0

ss

»0

41

4«

3%

10

If

20

3 4 5 6 9 KM 10» 11' 11" 12' 12» 1.» 1.11 2-7 2-1T J-M 3-1»

Fig. 6. Seasonal growth of male individuals in population of L. occipitalis, from March 1987 toMarch 1988.

«0

75

70

• 5

•o

55

50

45

40

35

30

25

20

1 4 5 • » » H I n* in i2» 1-î 1» *•» tn 3-" 3»

Fig. 7. Seasonal growth of female individuals in population of L. occipitalis, from March 1987 toMarch 1988.

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108 LAURA VERRASTRO AND LIGIA KRAUSE

Weight. Seasonal analysis of weight variation was carried out in the same fash-ion as the analysis of growth, with the purpose of studying the behavior of thisvariable in the development of the individuals.

During autumn, as with growth, there is a certain increase of weight for bothsexes, followed by a decrease during winter. It is during spring that individualsactually present a weight increase; it reaches maximum speed at the end of theseason, tending again to reduce its pace in summer (Figs. 8-9).

For Uta stansburiana, Tinkle (1967) found a similar seasonality in the behaviorof this variable. He explains the reduced weight during summer with the possiblyincreased activities of individuals as well as the higher environmental tempera-tures, when lizards would then spend more energy.

In relation to the females, this reduced weight increase during summer ismore evident because this is the season for laying eggs. Thus they present, asindividuals, a sudden decrease in weight during summer, which affects the gen-eral curve. Tinkle (1967), Gennaro (1974) and others mention this fact too.

Both males and females, throughout summer, continue gaining weight, thoughat a slower pace, a peak occurring at the end of the season. This was also ob-served for Uta stansburiana, by Tinkle (1997), and for Holbrookia m. maculata,by Gennaro (1974). These two authors explain that this phenomenon is bound tooccur because individuals are already building up fat in the pre-hibernal period.

If one carefully observes what happens with the females (Fig. 9), only adecrease in weight recurs (first two weeks of March). However, this deserves tobe investigated further and more accurately, for it could be explained either by a

» « » • » K» I M 1W 1W7 U-7 wi 17 V" 2' 2-17 Ï-I4 »:»DATE

19B7 „„

Fig. 8. Seasonal increase in weight of male individuals in population of L. occipitalis, from March1987 to March 1988.

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GROWTH IN A POPULATION OF LIOLAEMUS OCCIPTTAUS 109

12-

11

10

t

> 4 s e 101 »•« 11' 11" « ' 1*« M 111 I T 317 3i4 1 »

DATE

Fig. 9. Seasonal increase in weight of female individuals in population of L. occipitalis, from March1987 to March 1988.

characteristic of the females during this period or by deviations resulting fromthe few data obtained through collection in that period.

Finally, it was seen that both characteristics analysed, weight and SVL, aresimilar in terms of seasonality.

Conclusions

Two different aspects must be considered here:(1) Compared to other species of its genus, L. occipitalis seems to be, up to

now, the species that presents one of the smallest sizes among those speciesalready studied.

(2) Compared to other species of Tropiduridae and those of other families:(a) It was detected a direct relation between increase in weight and growth,

thus showing that L. occipitalis invests less in its growth than other species ofTropiduridae during juvenile stage;

(b) L. occipitalis follows the pattern in terms of growth rate, presenting amaximum at juvenile stage and decreasing as it reaches adulthood until it presentsvalues close to zero;

(c) A seasonality was established, similar to that presented by most lizards oftemperate zones, with high weight and growth rates during spring, rates thatcome close to zero during winter;

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110 LAURA VERRASTRO AND LIGIA KRAUSE

(d) Sexual dimorphism is well represented in the difference of size betweenmales and females, the latter being invariably smaller, just like most other liz-ards.

Taking into consideration that many species comparatively analysed hereinhabit regions of a more steady climate, without well-defined seasons (e.g. Utastansburiana, Tinkle, 1967 and Sceloporus occidentalis, Davis, 1968), it seemsto be quite clear that growth patterns follow the same general behavior of reptilesof temperate zones, to which the species Liolaemus occipitalis, object of thisstudy, fits into.

Acknowledgements

This study was sponsored by CAPES, through a Social Demand Scholarship for a Master's Degreefor the first author (1986-1989) and by CNPq, through a Research Grant (No. 000408825/87). We aregrateful for the invaluable help, in the field-work, from biologists Jairo Jose Zocche, Claudia Keller,and Angela M. Gallinati.

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