Habitat Preferences and Activity Patterns of Furcifer Pardalis

Habitat preferences and activity patterns of Furcifer pardalis

SALAMANDRA 44 3 129-140 Rheinbach, 20 August 2008 ISSN 0036-3375

Habitat preferences and activity patterns of Furcifer pardalis (Cuvier, 1829) in the Masoala Rain Forest Hall of the Zurich Zoo

Philip-Sebastian Gehring, Nicolà Lutzmann, Samuel Furrer & Roland Sossinka

Abstract. The habitat preferences and activity patterns of panther chameleons,Furcifer pardalis (Cuvier, 829) living in the Masoala Rain Forest Hall of the Zurich Zoo were investigated by means of radiotelemetry. Twelve animals were monitored over periods of up to 32 days. The animals exhibited a significant preference for the upper third, or crown stratum, of the plants. The chameleons were furthermore mostly encountered in the peripheral areas of their trees. Migratory patterns of male specimens in particular followed a largely linear course from the starting point. Distinctions could be made between specimens with a high degree of home range fidelity and more explorative individuals, which differed clearly with regard to the distances they moved away from their respective starting points. Key words. Reptilia, Chamaeleonidae, Furcifer pardalis, habitat preferences, activity patterns, telemetry, zoo.

Introduction concepts for the captive keeping of animals therefore have to be tested. The panther chameleon Furcifer ( pardalis) is a widely distributed and highly adaptable species of chameleon that is predominantly Material and methods found in anthropogenically altered and sec- The Masoala Rain Forest Hall ondary habitats (Ferguson et al. 2004, An- of the Zurich Zoo dreone et al. 2005, Lutzmann 2006). This gave rise to a number of studies on the ecol- The Masoala Rain Forest Hall of the Zurich ogy of F. pardalis (Bourgat 967, 968, 970, Zoo was opened in June of 2003. It measures Grimm & Ruckstuhl 999, Rimmele 999, 20 m in length, 90 m in width, and has a max- Ferguson et al. 2004, Andreone et al. 2005, imum height of 30 m (Fig. ). Thus providing Lutzmann 2006). With a floor space of ,000 a floor space of ,000 m², it is covered with m², the Masoala Rain Forest Hall of the Zu- some 7,000 plants, almost all of which are fit- rich Zoo presents an opportunity for studying ted with identification tags that not only give the behaviour of chameleons in a semi-natu- the respective species name, but also a numer- ral environment that about corresponds to ical ID number (IDN). These IDNs have been the natural habitats most commonly inhabit- plotted on a scaled map of the hall and were ed by F. pardalis in Madagascar. The manage- used to exactly pinpoint the locations of indi- ment of husbandry and captive breeding of vidual chameleons. animals in modern-day “eco-displays”, such as the Masoala Rain Forest Hall, necessitates that Radiotelemetry fundamental behavioural studies be conduct- ed in these artificial environments. These eco- Radiotelemetric monitoring was effected by displays replace some of the traditional exhibi- means of BD-2 transponders (Holohil Systems tion of animals in zoological gardens and new Ltd.) that have been tested with success in

felt pen on the front faces of their glued-on telemetry transponders. This enabled us to verify transponder numbers by means of bin- oculars. Permanent marking of the individuals was achieved by two other means. First, we counted the number of spikes in the individuals’ dorsal crests (crista dorsalis) from the head backwards and then removed one spike in a certain place. This method is entirely pain-free for the animals and has no eﬀect whatsoever on their behaviour, while it facilitates recogni- tion of individuals beyond doubt and serves as a lasting marker. Even though the scales constituting the dorsal crest grow with the animal, they will not regenerate fully, leaving a cropped tip visible. The second method involved the implantation of a passive integrat- ed transponder (PIT) manufactured by Trov- an (ID-00), which can be read by means of a portable scanner.

Fig. 1. View of the vegetation in the Masoala Rain Number of monitored specimens Forest Hall of Zurich Zoo. and monitoring previous studies of chameleons (Cuadrado The studied group comprised a total of twelve 2000, 200). These transponders have a range specimens, representing eleven males and one of about 200 m at a frequency of 48 kHz. A female. All animals were either just approach- portable R-000 Telemetry Receiver (Com- ing sexual maturity or had recently reached munications Specialists, Inc.) was used in con- this state. Table  shows the grouping of the in- junction with both an H- and a rod aerial. dividuals used. Transponders were fixed with medical Skin The chameleons were released into the Bond Cement, a wound glue manufactured hall in two groups. The first group (Group ) by Bruce Medical Supply. They were attached of five specimens was released on 24.0.2005, to the side of the dorsal crest above the hind with the second, comprising seven individu- limbs in all specimens participating in the als, following in two sub-groups (Groups 2/ first trial (Fig. 2), with the aerial being aligned and 2/2) on 28.02.2005. The two groups were parallel to the tail so that it would not affect released in different spots. While Group  was movement of either the tail or the hind limbs. released in the western section of the hall, near The second trial made use of three random- the public entrance, Group 2/ was set free in ly selected specimens, which were fitted with the southern sector, which is an area not acces- transponders on the side of the dorsal crest sible to the public and has the highest degree above the front limbs. of sun-exposure. Group 2/2 was released in the northern sector, which is likewise not open to the public, but is marked by less sun-exposure Marking of individuals and higher vegetation. The variety of release points was chosen to investigate the possible The participating individuals were marked influences of environmental factors on the be- simply by writing individual numbers with a haviour of the animals.

130 Habitat preferences and activity patterns of Furcifer pardalis

The animals were telemetry-located four times every day during the course of the study period, at intervals of one to two hours during the day. The times at which they were traced were varied so that a record would be available for every specimen for every hour within a time span of at maximum ﬁve days. Data col- lection was limited to the time between sunrise and sunset. With chameleons being diurnal, they were not expected to relocate at night.

Recorded data

Once a chameleon had been located, the time and the exact location were recorded. The IDN of the respective tree or shrub was noted. The animal’s position in the plant was evaluated according to standard categories, i.e., whether it was on the ground, or perched on the trunk, a branch, or a leaf. A tree was stratiﬁed into lower third, midriﬀ, and crown. Peripheral perch sites were recorded as such. The height of the perch above the ground was estimated in metres, more readily accessible perches were measured in centimetres. Fig. 2. Male Furcifer pardalis carrying a glued-on BD-2 transponder. Data analysis

The data gathered thus were analysed using to a level of significance of α = 0.05 at maxi- the software Statistika. The tests were limited mum. Horizontal migrations were calculated by means of the GIS software ArcView 3.2. Tab. 1. Distribution of specimens and their sexes The individual points of sightings were plot- in three groups. ted from the individual points of release to the Group Specimen No. Sex last sighting, graphically connected by straight lines, and these were then measured and to-  A ,0 talled. The distances between two points were  A2 ,0 identified by measuring them on a scaled map  A3 ,0 of the hall.  A4 ,0 Perch heights (h / h2) were calculated on  A5 0, the basis of the Pythagorean theorem, with 2/ B ,0 the horizontal distance between two perches 2/ B2 ,0 forming the first short side of a rectangular tri- 2/ B3 ,0 angle (d) and the difference between the two 2/ B4 ,0 perch heights (h - h2) the second. The long 2/2 C ,0 side (hypotenuse) of the resultant rectangular 2/2 C2 ,0 triangle thus equals the calculated distance of 2/2 C3 ,0 relocation of the animal:

131 Philip-Sebastian Gehring et al.

SitzhöhenPerch heights von Gruppe of group 1 1 8

7 7,00m 6

3 2,75m 2,50m 2,50m 2,00m Perch height Perch (m) 2 Sitzhöhe (m)

0 Median 25%-75% -1 A5 A4 A3 A2 A1 Min-Max IndividualTier specimen Numme numberr Fig. 3. Perch heights of the specimens in group 1. Box and whisker plots: The box comprises percentages 25-75% of the observations, while the open squares indicate means. The whiskers delimit minima and maxima. The Y-axis shows perch heights in metres, while the X-axis shows the individual specimens. Observations: A5 n=78; A4 n=120; A3 n=35; A2 n=15; A1 n=63. Standard deviations: A5 - 1.34 m; A4 - 1.02 m; A3 - 0.83 m; A2 - 1.87 m; A1 - 1.66 m.

species of plants or trees present in the hall. Quite to the contrary, they even made use of plants (e.g., Pandanus baptiste, Bismarckia no- Because it may be supposed that the ani- bilis, Ravenala madagascariensis) and struc- mals migrated from one tree to another via tures (e.g., hot air pipes and ventilation grilles) neighbouring branches rather than crossing that would appear rather unsuited for chame- over on the ground, especially in densely vege- leons. Water surfaces were crossed via the lush tated areas, this method of calculation appears vegetation of aquatic plants (Pistia stratiotes). to be fairly realistic. A total of 505 perch heights were recorded

Within a tree, the vertical migration (av) during the monitoring period. Figure 3 illus- was calculated as the difference (av = h - h2) trates the perch heights recorded for the ani- between two perch heights. mals in Group . The mean value for the perch The extent of activity was always evaluated heights of the five participating individu- for the last eight days of the study period of als amounts to 3.29 m (n = 3; SD = .34 m; the respective specimen. min. = 0 m, max. = 7 m). The mean of perch heights recorded for the individual animals lies between min. = 2,00 m and max. = 7,00 Results m. Half of all observations involved animals Habitat utilization that were perched at heights above two metres. The three groups differed substantially Our observations did not suggest to animals with regard to the distribution of their perch to have any particular predilection for certain heights. Figure 4 illustrates the distribution of

132 Habitat preferences and activity patterns of Furcifer pardalis

SitzhöhenPerch heights von Gruppeof group 2/1 6

2,00m 2,00m 2

1,80m Perch height Perch (m) 1 1,17m

Median -1 B1 B2 B3 B4 25%-75% Min-Max IndividualTier specimen Numme numberr Fig. 4. Perch heights of the specimens in group 2/1. Box and whisker plots: The box comprises percentages 25-75% of the observations, while the open squares indicate means. The whiskers delimit minima and maxima. The Y-axis shows perch heights in metres, while the X-axis shows the individual specimens. Observations: B1 n=30; B2 n=46; B3 n=10; B4 n=11. Standard deviations: B1 - 0.64 m; B2 - 1.02 m; B3 - 0.50 m; B4 - 0.50 m.

SitzhöhenPerch heights von ofGruppe group 2/2/22 8

5,00m 5

3,00m

Perch height Perch (m) 3 2,50m

2 Sitzhöhe (m)

Median 0 C1 C2 C3 25%-75% Min-Max IndividualTi erspecimen Numme numberr Fig. 5. Perch heights of the specimens in group 2/2. Box and whisker plots: The box comprises percentages 25-75% of the observations, while the open squares indicate means. The whiskers delimit minima and maxima. The Y-axis shows perch heights in metres, while the X-axis shows the individual specimens. Ob- servations: C1 n=21; C2 n=41; C3 n=35. Standard deviations: C1 - 2.03 m; C2 - 1.60 m; C3 - 1.52 m.

133 Philip-Sebastian Gehring et al. Verteilung peripherer und nicht- peripherer Sitzplätze

13,4% values likewise refer only to the last twenty observations. 46,7% 53,3% non-Nicht – peripheralperipherer peripheralPeripherer perch perchSitzplatz Sitzplatz 86,6% Activity patterns and spatial utilization

During the observation period, the individual

26,2% Von links animals relocated over varying distances from nach rechts: the original point of release. Table 2 summa- Gruppe 1 FromGuppe left 2/ 1to right: rizes these relocations of the individual ani- GroupGruppe 1 2/2 Group 2/1 mals. This is based merely on a supposed two- 73,8% Group 2/2 dimensional distance between the point of Fig. 6. Illustration of the distribution of peripheral release and the location of the last recording, and non-peripheral perches within the utilized however (see 2.6). plants for the individual groups. The peripheral Displacing itself by a total of 29.50 m with- proportions are shown as red solid areas and in 2 days, a male from Group 2/ (B) trav- the non-peripheral proportions in green dotted elled the greatest distance. A male from Group areas. 2/2 (C3), on the other hand, covered only a distance of 2.40 m within a similar period perch heights noted for Group 2/. This group of time (4 days). In general, it is obvious that produced a mean perch height of .88 m (n= the animals in Group 2/ walked substantially 97; SD = 0.89 m; min.: 0 m, max.: 5 m). More larger distances during the observation period. than 50% of the observations involved animals Every animal in this group covered a distance perched at heights between one and three me- of about 00 m within 4 days, whereas these tres. The perch heights recorded for Group 2/2 distances averaged only about 40 m (Group ) are higher than those for Group 2/. The mean and 20 m (Group 2/2) within 4 days, respec- value for the perch heights recorded for Group tively. In order to create a nearly three-dimen- 2/2 lies at 3.54 m (n = 97; SD = .7 m; min.: sional impression of the distances really cov- 0.30 m, max.: 7.00 m). Figure 5 is a graphic il- ered by the animals, their activity ranges (a) lustration of the distribution of perch heights were calculated in addition to the individually of the individual animals of Group 2/2. recorded perch heights (h) (see 2.6). Further-

A comparison of the utilized tree strata more, all vertical relocations (av) within a tree showed that the perches recorded from within that were not included in the activity range a tree highly significantly deviated from a bal- were worked out. The activity ranges were anced utilization of all strata available within a evaluated for the last eight days of the obser- tree (χ2 -Test; p < 0,0). All observations of all vation period for each animal. The results are animals instead revealed a significant predilec- presented in Table 3. tion for the upper third, respectively the crown The graphic illustration of the distances stratum (H-Test; p < 0.0; mean: 3.0) of plants. covered by the chameleons (Fig. 7) reveals The three groups showed differences, however. two migratory patterns, which are distributed Both Groups  and 2/ had a significant predi- unevenly between the three groups. Group  lection for the top third respectively the crown and 2/2 both comprised animals with relative- stratum (H-Test; p < 0.0; mean: 3.0). Group 2/2 instead exhibited a significant predilection right page: Fig. 7. Illustration of distances covered for the middle third (H-Test; p < 0.0; mean by the individual specimens. The recorded perch 2.0). Figure 6 illustrates that the animals of all sites are marked with symbols whose sizes are three groups were relatively more often spot- correlated to the numbers of days on which the ted in peripheral regions of their trees. These animals were found in these spots.

134 Habitat preferences and activity patterns of Furcifer pardalis

135 Philip-Sebastian Gehring et al.

Tab. 2. Overview of distances travelled by individual specimens and their groups from the original point of release to the last sighting. Distances are given in metres and refer only to the assumed two-dimensional displacement. The observation period is given in days.

A A3 A4 A5 B B2 B3 B4 C C2 C3 Group     2/ 2/ 2/ 2/ 2/2 2/2 2/2 Distance in m 2.30 6.80 80.30 25.60 29.50 94.0 49.20 08.90 8.50 55.0 2.40 Period in days 5 4 29 3 2 5 8 4 5 5 4 ly high degrees of site fidelity and those that Regular shedding enabled all animals to rid were more explorative. In Group , the animals themselves of remains of the glue, not leaving A and A5 presented themselves as rather sta- any residue or causing skin irritation. tionary, and in Group 2/2, animal C3 was notable for displacing itself only by a small distance from the start-off point. The members Discussion of Group 2/ all covered larger distances from Choice of habitat their point of release. The activity patterns show a distinctly linear course, especially as The panther chameleons living in the Masoa- far as Group 2/ is concerned. la Rain Forest Hall exhibited a significant predilection for the upper third of the plants utilized. Living mainly in an area with numer- Durability of transponder attachment ous large trees (> 4 m), the animals of Group  were particularly commonly found in the top The period of time throughout which the ani- third or crown stratum. All groups exhibited a mals carried their transponders, and the du- predilection for the middle and top thirds of rability of the attachment (glue) in particu- their plants. The animals were encountered in lar, differed substantially between the three the bottom third of their plants rather rarely groups. The animals of the first group carried and usually only when they were on the move. their transponders for an average of 8 days Furthermore, the panther chameleons appar- and could each be observed for another four ently spent more time in the peripheral areas days on average after they had lost their trans- of their plants than elsewhere. These results ponders. suggest that panther chameleons will tend to In the second group, transponders re- select perches at greater heights in an environ- mained affixed for only 6.5 days on average, ment that offers tall trees and there preferably which equals half of the average total observa- utilize the peripheral areas. This confirms the tion period of 3 days. Some specimens could observations made by Glaw & Vences (994) be regularly traced even after they had lost and Raxworthy (988). Because the availa- their transponders also here. Female A5 was bility of microclimates that can facilitate ther- the animal observed longest, i.e., for a total of moregulation is one of the most important de- 3 days. Specimens A carried its transponder terminants for the choice of habitat in reptiles for the longest period of time, i.e., for 32 days. (e.g., Zug et al. 200), higher ambient tem- The two different points of attachment peratures and the availability of basking spots (above the front limbs vs. above the hind could offer an explanation for a predilection limbs) proved to be varyingly well suited, with for the crown stratum. Research on the dis- all three animals on which the transponder tribution of microclimates in rain forests has had been affixed above the shoulder losing shown that temperatures rise higher during their equipment within the first three days af- the day in the upper storeys of trees than on ter release. the forest floor (comp., e.g., Terborgh 993).

136 Habitat preferences and activity patterns of Furcifer pardalis

Panther chameleons inhabit within their dis- by studies of the top storey of the Masoala rain tribution range a wide variety of vegetation forest by means of an airship, which produced types and landscapes, rendering descriptions records of only one species of chameleon, i.e., F. of natural habitats partly highly contradic- pardalis (Andreone et al. 2005), even though tory. They mention in common, though, a it remained unclear whether this undertaking presence of trees, bushes and shrubs in open was in search of chameleons or how they were landscapes and cultivated areas with low sec- traced. However, futher studies in the canopy ondary vegetation, including human settle- of malagasy rainforests need to be carried out, ments (e.g., Andreone et al. 2005, Ferguson to clarify the occurrence and the densities of F. et al. 2004, Glaw & Vences 994, Henkel pardalis in the forest overstory. F. pardalis was & Heinecke 993, Henkel & Schmidt 995, most commonly encountered in secondary Müller et al. 2004, Necas 2004, Schmidt et vegetation or anthropogenously altered habi- al. 996, own obs.). However, descriptions of tats. In forested areas, F. pardalis was mostly primary rain forest habitats in which F. parda- found along rivers or in clearings, even though lis occurs differ substantially: while some au- Lutzmann (2006) observed females laying thors explicitly exclude dense forests (Hen- their eggs in closed forest areas. Andreone et kel & Heinecke 993, Henkel & Schmidt al. (2005) presumed that F. pardalis preferred 995, Schmidt et al. 996, Schmidt & Liebel these open areas because unimpeded inso- 2004), others have published records of F. par- lation would create optimum conditions for dalis from rain forest regions (Andreone et basking and visual communication. al. 2000, 2005, Glaw & Vences 994, Klin- According to this, the panther chameleon gelhöfer 957, Lutzmann 2006, Müller et appears to have been a “forest edge species” al. 2004, Ramanantsoa 974, Raxworthy originally, which lived in open spaces and mar- 988, Rimmele 999, own obs.). It is notable, ginal zones of the once-expansive rain forests. though, that population densities of F. parda- As the destruction of the forests progressed, lis, as derived from transect studies, are much more and more open areas and cultivated lower in rain forest regions than in areas with land with low secondary vegetation emerged, secondary vegetation, even when the respec- which appear to offer panther chameleons op- tive authors tried to also search the crowns of timum conditions. Consequently, high popu- trees from the ground (Andreone et al. 2005). lation densities are found in exactly these areas Raxworthy (988), Glaw & Vences (994), (Andreone et al. 2005, Lutzmann 2006). and Lutzmann (2006) presumed that F. pard- The preferred utilization of the periph- alis would be exclusive to the crown stratum in eral areas of trees could also be a function of primary rain forests because only this region the more intense insolation in these parts of would permit sufficient insolation to satisfy the trees. Also imaginable would be an anti- the thermal requirements of these distinctly predator strategy, as potential, larger preda- heliophilic chameleons. This is also indicated tors would be unable to follow the chamele-

Tab. 3. Overview of individual activity ranges (a) and vertical displacement (av) of the individual chameleons during the last eight days of observation. * = Specimens for which no continuous data were available for the last eight days (A1: 5 days; A3: 4 days; B1: 6 days).

A A3 A4 A5 B B2 B3 B4 C C2 C3 Group     2/ 2/ 2/ 2/ 2/2 2/2 2/2 a (m) 4.00* 4.40* 33.50 5.00 50.60* 5.30 46.50 57.30 9.83 33.0 .20 av (m) 3.50 4.50 5.40 2.0 2.00 6.80 .00 29.00 8.50 5.70 5.00

∑a (m) 7.50 28.90 38.90 7.0 62.60 58.0 47.50 86.30 28.33 48.80 26.20

137 Philip-Sebastian Gehring et al. ons out onto the thinner peripheral branch- son et al. (2004) even noted it in some males es, or would at least announce their presence during the mating season. through vibrations (“perch release”, Parcher The activity patterns displayed by males 974). As a result of the highly biased sex ratio (Fig. 5) followed a highly linear course. The in our study groups, no statistical evaluation young males of Group 2 covered particularly of a possibly sex-dependent choice of habitat long distances. This might suggest thatF. par- could be performed. Andreone et al. (2005) dalis males do not establish permanent “terri- noted a significant difference at least with re- tories”, but rather roam their environment in gard to perch heights between male and fe- search of food, or, during the mating season, male F. pardalis living on Nosy Be, which was in a quest to locate a female. If the environ- explained by a higher perch providing males mental conditions turn unfavourable (tem- with a better overview of their territories. peratures, weather or others), they will stay in Lutzmann (2006) did not find differences in an area for some time without displaying ter- perch heights between males and females in ritorial behaviour towards other males. The Maroantsetra, but could also not detect indi- observations made in the Hall, as well as dur- cations of male territoriality. ing studies of Chamaeleo chamaeleon (Cuad- rado 200), suggest that females maintain fixed home ranges instead. If a male happens Activity patterns and spatial utilization to come upon a female, he will stay in her vicinity for some days and mate with her. In an As Figure 7 illustrates, it showed that there attempt to secure his own reproductive suc- were two types of activity patterns present: the cess, he will remain in the female’s immediate more resident type and the more explorative vicinity and defend her with a great degree type. Both types were notable for their typi- of aggression against potential competitors. cally linear movements from a start-off point, Whether he will copulate more than once i.e, both rarely visited the same area more than with this particular female during this period once. Specimens A5, A and C3 presented of time is as yet unknown. “Female-guard- themselves as rather stationary and this was ing” was also noted by Lutzmann (2002) particularly evident in the female A5. While in the case of Chamaeleo africanus in Egypt. this animal stayed within a certain range “Partner-bonding” in F. pardalis during the throughout the entire observation period, the mating season was reported about by Bour- males A and C3 showed a linear type of re- gat (970) and found confirmation in obser- location. Even though not as expressed as in vations made following a successful mating other specimens, female A5 moved about con- observed in the Masoala Rain Forest Hall tinually within a certain perimeter and visit- during the present study. Whether the males ed certain spots repeatedly. She also relocated will resume their meandering after a while between various perches during the course in order to find and mate with other females of a day. During phases of sexual inactiv- has not been proven, but it appears likely. ity, which in the wild coincides with the dry Mated females exhibit very strong defensive season (Müller et al. 2004, Andreone et al. responses toward other males. A polygynic 2005), inter-male competition is not particu- mating system of this kind has also been not- larly expressed. This must surely be viewed as ed in the case of C. chamaeleon (Cuadrado an avoidance of energy-intensive squabbles at 200). The males would guard their females a time when resources are limited. In the Hall, for a number of days after copulation, until males were repeatedly seen right next to each the latter clearly showed by means of their other without showing any gestures of aggres- colour pattern that they would not be recep- sion. This peaceful coexistence was also de- tive to a male any longer. The guarding be- scribed by Müller et al. (2004), and Fergu- haviour of the males culminates in a strongly

138 Habitat preferences and activity patterns of Furcifer pardalis territorial display within the home range of History, Conservation, and Captive Manage- the female in that it is defended against po- ment. – Krieger Publishing Company, Malabar, tential competitors. Females, on the other Florida, pp. 25. hand, do not exhibit competitive behaviour Glaw, F. & M. Vences (994): A Fieldguide to towards other females. the Amphibians and Reptiles of Madagascar. – 2nd edition including mammals and freshwa- ter fish. – M. Vences & F. Glaw Verlags – GbR, Acknowledgements Köln, 480 pp. Henkel, F. W. & S. Heinecke (996): Chamäleons We thank Zurich Zoo for the opportunity to carry im Terrarium. (2nd edition) – Landbuch-Ver- out this study in their Masoala Rain Forest Hall lag, Hannover, 58 pp. and for making available telemetry equipment. Henkel, F. W. & W. Schmidt (995): Amphibien Many thanks go to the caretakers and the team of und Reptilien Madagaskars, der Maskarenen, volunteers of Zurich Zoo for their assistance dur- Seychellen und Komoren. – Ulmer, Stuttgart, ing the gathering of data. 3 pp. Klingelhöfer, W. (957): Terrarienkunde, 3. Teil References Echsen. – A. Kernen Verlag, 246 pp. Lutzmann, N. (2002): Untersuchungen zur Ver- Andreone, F., F. M. Guarino & J. E. Randri- breitung, Systematik und Ökologie der Cha- anirina (2005): Life history traits and age pro- mäleonfauna Ägyptens. – Thesis University of file as useful conservation tools for the panther Bonn, unpubl. chameleons (Furcifer pardalis) at Nosy Be, NW Lutzmann, N. (2006): Untersuchungen zur Öko- Madagascar. – Tropical Zoology, 8: 209-225. logie der Chamäleonfauna des Nationalparks Andreone, F., J. E. Randrianirina, P. D. Jenkins Masoala in Nordost Madagaskar. – Dissertati- & G. Aprea (2000): Species diversity of Am- on University of Bonn, unpubl. phibia, Reptilia and Lipotyphla (Mammalia) Müller, R., N. Lutzmann & U. Walbröl (2004): at Ambolokopatrika, a rainforest between the Furcifer pardalis – Das Pantherchamäleon. Anjanaharibe – Sud an Marojejy massifs, NE – Natur und Tier-Verlag GmbH, Münster, 27 Madagascar. – Biodiversity and Conservation pp. 9: 587-622 Nečas, P. (2004): Chamäleons – Bunte Juwe- Bourgat, R. M. (968): Comportment de la fe- len der Natur (3. Aufl.). – Edition Chimaira, melle de Chamaeleo pardalis Cuvier 829 Frankfurt a. M., 382 pp. après láccouplement. – Bull. Soc. Zool. Fran- Parcher, S. R. (974): Observations on the Natu- ce, 93: 355-356. ral Histories of Six Malagasy Chamaeleontidae. Bourgat, R. M. (970): Recheres écologiques sur – Zeitschrift für Tierpsychologie 34: 500-523. le Chamaeleo pardalis Cuvier 829 de I´île Ramanantsoa, G.A. (974): Connaissance des Réunion et Madagascar. – Bull. Soc. Zool. Caméléonides communs de la Province de France, 95: 259 – 269. Diégo-Suarez par la Population Paysanne. – Cuadrado, M. (2000): Body Colors Indicate the Bull. Acad. Malg., 5(): 47-49. Reproductive Status of Female Common Cha- Raxworthy, C. J. (988): Reptiles, rainforest and meleons: Experimental Evidence for the Inter- conservation in Madagascar. – Biological Con- sex Communication Function. – Ethology 06: servation, 43: 8-2. 79-9. Rimmele, A. (999): Vorstellung der in der Zucht- Cuadrado, M. (200): Mate guarding and social gemeinschaft Chamaeleonidae gezüchteten mating system in male common chameleons Chamäleons – Teil VI: Erkenntnisse aus der (Chamaeleo chameleon). – J. Zool. London, mehrjährigen Pflege und Zucht, sowie einige 255: 425-435. Freilandbeobachtungen am Pantherchamäle- Ferguson, G. W., J. B. Murphy, J.-B. Ramana- on, Furcifer pardalis (Cuvier, 829). – Sauria manjato & A. P. Raselimanana (2004): The 2: 27-36. Panther Chameleon – Color Variation, Natural

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Schmidt, W., K. Tamm & E. Wallikewitz (2000): Terborgh, J. (993): Lebensraum Regenwald: Chamäleons – Drachen unserer Zeit (2nd edi- Zentrum der biologischen Vielfalt. – Spektrum tion). – Natur- und Tier-Verlag, Münster, 60 Akademischer Verlag, Heidelberg, Berlin, New pp. York, 253 pp. Schmidt, W. & K. Liebel (2004): Furcifer: Cha- Zug, G. R., L. J. Vitt & J. P. Cladwell (200): mäleons aus Madagaskar. – DATZ 57: 36-42. Herpetology – An Indtroductory Biology of Amphibians and Reptiles (2nd Edition). – Aca- demic Press, San Diego, London; 640 pp.

Manuscript received: 18 March 2007 Authors’ addresses: Philip-Sebastian Gehring, Grewenbrink 5a, D-33619 Bielefeld, Germany, E-Mail: [email protected]; Nicolà Lutzmann, Zoologisches Forschungsmuseum A. Koenig, Ade- nauerallee 160, D-53113 Bonn, Germany, E-Mail: [email protected]; Samuel Furrer, Zoo Zurich, Zurichbergstrasse 221, CH-8044 Zurich, Switzerland, E-Mail: [email protected]; Roland Sossinka, Department of Animal Behavior, University of Bielefeld, Morgenbreede 45, D-33615 Bielefeld, Germany, E-Mail: [email protected].

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