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Comparative Ecology, Resource Utilization and Niche Segregation Among Gekkonid Lizards in the Southern Kalahari

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Comparative Ecology, Resource Utilization and Niche Segregation Among Gekkonid Lizards in the Southern Kalahari JAEGER-SALAMANDER PLANT CLIMBING 691 ACKNOWLEDGMENTS JAEGER,R. G. 1972. Food as a limited resource in competition between two speciesof terrestrialsala- Research was supported by American Philo- manders. Ecology 53:535-546. sophical Society grant 7113, by the Research . 1974. Competitive exclusion: comments Foundation of the State of New York on survival and extinction of species. BioScience University 24:33-39. 20-7331-A and funds from the Shenan- grant by KIESTER,A. R., ANDM. SLATKIN. 1974. A strategy doah Natural History Association. I am very of movement and resource utilization. Theor. grateful to Linda Jaeger and William Gergits for Pop. Biol. 6:1-20. in the field studies and to L. for MADISON,D. M. 1972. Homing orientation in sala- help Jaeger manders: a mechanism the involved. involving chemical cues, computer programming Douglas p. 485-498. In: Animal orientation and naviga- Fraser, W. Gergits and Patricia Nanartowich tion. S. R. Galler, K. Schmidt-Koenig, G. J. Jacobs kindly criticized the manuscript. I thank E. and R. E. Belleville (eds.). Nat. Aeronautics and Raymond Schaffner and Dennis Carter, of the Space Admin., Washington, D.C. SAMUELSEN,P. 1977. Plethodon cinereus home National Park Service, for to work permission range and homing. Unpubl. M.S. thesis, Univ. in Shenandoah National Park. Rhode Island. SIEGEL,S. 1956. Nonparametric statistics for the LITERATURECITED behavioral sciences. McGraw-Hill Book Co., New York. FRASER,D. F. 1976. Empirical evaluation of the U.S. DEPARTMENTOF COMMERCE.1970. Climato- hypothesis of food competition in salamandersof logical Data 80: nos. 7 and 8. the genus Plethodon. Ecology 57:459-471. GIST, C. S., AND D. A. CROSSLEY.1975. The litter DEPARTMENT OF BIOLOGICAL SCIENCES, STATE arthropod community in a southern Appalachian hardwood forest: numbers, biomass and mineral UNIVERSITYOF NEW YORK,ALBANY, NEW YORK element content. Amer. Midi. Natur. 93:107-122. 12222. Accepted 26 Feb. 1978. Copeia, 1978(4), pp. 691-701 Comparative Ecology, Resource Utilization and Niche Segregation Among Gekkonid Lizards in the Southern Kalahari ERIC R. PIANKA AND RAYMONDB. HUEY Varied aspects of the general ecology of little-known gekkonid lizards from the southern Kalahari semi-desert of Africa are documented. Four to seven species of geckos coexist on ten study areas. Six of the seven species are nocturnal: three species are ground-dwellers, three climb (of these, one is diurnal) and one is semiarboreal. The percentage of climbing species in various areas is positively correlated with plant height diversity. Body tem- peratures of nocturnal geckos are very similar and these lizards thermo- regulate less carefully than diurnal Kalahari lizards. Pairs of species differ in habitat, microhabitat and/or diet. Resource utilization patterns among Kalahari geckos are compared with those of an independently-evolved, but otherwise ecologically similar, nocturnal saurofauna: the geckos of the western Australian desert. Microhabitat utilization is similar on both con- tinents, but Australian geckos eat a greater variety of prey taxa than Kalahari geckos. Pairs of morphologically similar species on the two continents do not necessarily converge ecologically in diet and micro- habitat. The frequency of pairs with high overlap in both diet and microhabitat is greater among intercontinental comparisons than it is among intracontinental ones, suggesting a limit to the similarity of poten- tially competing species. GECKOS are an important component of Africa, geckos comprise from 31 % to 44% (mean many desert lizard faunas, particularly in 36%) of the total saurofauna on ten selected Africa and Australia (Pianka, 1973, 1975). In study areas differing in topography, vegetation, the southern Kalahari semi-desert of southern and climate (Pianka, 1971; Pianka and Huey, ? 1978 by the American Society of Ichthyologists and Herpetologists 692 COPEIA, 1978, NO. 4 similar data on geckos from an independently- B evolved, yet ecologically very similar, nocturnal - . 50 saurofauna: the geckos of the Great Victoria Desert of Western Australia (Pianka and Pianka, x L K * * i 1976). Our approach is somewhat eclectic but 40 - fill a of LJ intentionally designed to help variety in knowledge about the general ecology of o gaps cQ MDA nocturnal cr - S** geckos. 30 Elsewhere we have evaluated overall niche z- of these in the context of z relationships geckos LJ of entire saurofaunas (Pianka, 1973, 20 - analyses U 1974, 1975; Pianka, Huey and Lawlor, 1978), LL-LU data on the se. G_ -G R without presenting geckos per * described 10 Haacke (1975, 1976a,b) recently systematics and briefly discussed aspects of the natural history of the three ground-dwelling liz- and 0 ards (Colopus, Chondrodactylus Ptenopus). .2 .4 .6 .8 1.0 Leistner (1967) described the vegetation, top- the Kalahari in PLANT HEIGHT DIVERSITY ography and climate of southern detail. Stabilized sandridges cover most of the extensive Fig. 1. Plot of the percentageof gecko species that southwestern region and grade into was arboreal versus plant height diversity for ten areas of sandplain to the east. Vegetation is pre- study areas in the Kalahari (r, = .59, P < .05). somewhat savanna-like, with Letters sites as listed in dominantly grassy, designate particular study scattered shrubs and some small trees (im- Pianka (1971). many portant to climbing geckos) including Acacia low A. 1971). During the course of a year-long field mellifera (a sprawling shrubby plant), tree with dense investigation of the ecology and diversity of haemotoxylon (a small foliage), bark and lizards on these study areas, we gathered eco- A. giraffae (a large tree with rough logical data on most of the seven species of dense foliage) and Boscia albitrunca (a moderate- previously little-known but widespread geckos. sized tree with relatively smooth pale bark and Our data provide important new information hollows). Fallen logs are common in some areas. on the comparative ecology, reproduction, broken tail frequencies, thermal relations, diet, METHODS habitat and microhabitat associations, and niche were segregation of these lizards. Moreover, these Active nocturnal geckos spotted by eye shine head We re- data permit informative comparisons with shine or body using lamps. and TAPLE 1. OCCURRENCES OF DIFFERENT GECKO SPECIES ON THE VARIOUS STUDY AREAS. See Fig. 1 in Huey Pianka (1974) for localities coded by letters. L K M B A X G D R Ts Tf T Microhabitat Retreat x x terrestrial burrow Colopus wahlbergi x x x x x x x x e x burrow Chondrodactylus angulifer x x x x x x x x x x x x terrestrial burrow Ptenopus garrulus x x x x x x x x x x x x terrestrial termite mounds Pachydactylus capensis x e x x x x x x x x x x semiarboreal under bark, log xxx arboreal under and in Pachydactylus rugosus xxx x log or bunch grass x under bark, Pachydactylus bibroni xx x x x arboreal l3gs, rocks under Lygodactylus capensis x arboreal bark, logs x = collected, e = highly expected. PIANKA AND HUEY-KALAHARI GEKKONID ECOLOGY 693 TABLE 2. MICROGEOGRAPHICALASSOCIATIONS OF GECKOSIN SANDRIDGELOCALITIES (PERCENTAGES). Chondro- Colopus dactylus Ptenopus P. capensis P. rugosus P. bibroni Lygodactylus Flat 12.8 70.9 79.2 93.8 22.2 83.0 96.3 Base 12.8 7.8 9.4 6.3 11.1 5.3 0 Slope 59.0 8.6 7.5 0.0 61.1 8.5 0 Crest 15.4 12.7 3.8 0.0 5.6 3.2 3.7 N 39 268 106 16 18 94 27 corded microhabitat, habitat and time of ac- is arboreal is significantly correlated with plant tivity, as well as body (cloacal) and air tem- height diversity (rs = .71, P < .05; Fig. 1). peratures at capture with Schultheis thin-bulb Thus areas with high vegetative diversity support thermometers. To obtain an indication of the more species of arboreal geckos, a pattern that general ambient thermal conditions, air tem- holds as well for numbers of bird species (Pianka peratures were recorded at chest height rather and Huey, 1971) and for number of species of than at the point of capture (other studies have diurnal lizards in the Kalahari (rs = .71, P < shown that body temperatures correlate better .05). with chest height air than with temperatures distributions on sites. ground-level air temperatures, e.g., Schall, 1977). Microgeographic sandridge -We four subdivi- Techniques of analysis of reproductive traits recognize microgeographic sions of dunes and Pianka, in sand- and stomach contents using preserved material (Huey 1974) areas: street, base, and crest parallel those of Pianka and Pianka (1976). ridge slope, (Table 2). While most species are found primarily in dune streets, do differ RESULTS geckos significantly (G- test, P < .001) in microgeographic distributions. Occurrences on study sites.-Gecko censuses on However, Colopus and P. rugosus (both ex- ten study areas, microhabitat associations while tremely rare geckos before our study), which active, and diurnal retreat of each species are themselves do not differ significantly (STP-test, presented in Table 1. The number of sympatric P > .9), are primarily associated with dune gecko species varies from 4 to 7 and is signifi- slopes (they occasionally occur in flats, too). cantly correlated with plant height diversity at [The association of P. rugosus with sandridge the site (rs = .59, P < .05; plant data in Pianka, slopes may reflect the frequent occurrence of 1971). this species on sprawling Acacia shrubs (especi- The three ground-dwelling species are nearly ally A. mellifera but also A. haemotoxylon, see ubiquitous, but the occurrence of climbing below) that are most common on dune slopes species is spotty (Table 1). This distributional (Leistner, 1967).] Similar differences in micro- distributions occur in Kalahari sub- difference presumably reflects the greater sensi- geographic terranean lizards and Pianka, and tivity of arboreal species to vegetation structure. (Huey 1974) in Australian lizards 1969, 1972, The percentage of gecko species on an area that (Pianka, 1978).
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