Natural restoration of the species-area relation for a lizard after a hurricane.

We document the decimation and recovery of the commonest lizard species, Anolis sagrei, on 66 islands in the Bahamas that were directly hit by Hurricane Floyd in September 1999. Before the hurricane, an island's area was a better predictor of the occurrence of A. sagrei than was its altitude. Immediately after, altitude was a better predictor: Apparently all lizards on islands lower than about 3 meters maximum elevation perished in the storm surge. After about 1 year, area again became the better predictor. By 19 months after the hurricane, A. sagrei populations occurred on 88% of the islands they formerly occupied. Recovery occurred via overwater colonization and propagation from eggs that survived inundation, mechanisms that were enhanced by larger island area. Thus, natural processes first destroyed and then quickly restored a highly regular species-area distribution.

Predators increase the risk of catastrophic extinction of prey populations.

There has been considerable research on both top-down effects and on disturbances in ecological communities; however, the interaction between the two, when the disturbance is catastrophic, has rarely been examined. Predators may increase the probability of prey extinction resulting from a catastrophic disturbance both by reducing prey population size and by changing ecological traits of prey individuals such as habitat characteristics in a way that increases the vulnerability of prey species to extinction. We show that a major hurricane in the Bahamas led to the extinction of lizard populations on most islands onto which a predator had been experimentally introduced, whereas no populations became extinct on control islands. Before the hurricane, the predator had reduced prey populations to about half of those on control islands. Two months after the hurricane, we found only recently hatched individuals--apparently lizards survived the inundating storm surge only as eggs. On predator-introduction islands, those hatchling populations were a smaller fraction of pre-hurricane populations than on control islands. Egg survival allowed rapid recovery of prey populations to pre-hurricane levels on all control islands but on only a third of predator-introduction islands--the other two-thirds lost their prey populations. Thus climatic disturbance compounded by predation brought prey populations to extinction.

Experimental studies of adaptive differentiation in Bahamian Anolis lizards.

Populations of the lizards Anolis carolinensis and A. sagrei were experimentally introduced onto small islands in the Bahamas. Less than 15 years after introduction, we investigated whether the populations had diverged and, if so, whether differentiation was related to island vegetational characteristics or propagule size. No effect of founding population size was evident, but differentiation of A. sagrei appears to have been adaptive, a direct relationship existed between how vegetationally different an experimental island was from the source island and how much the experimental population on that island had diverged morphologically. Populations of A. carolinensis had also diverged, but were too few for quantitative comparisons. A parallel exists between the divergence of experimental populations of A. sagrei and the adaptive radiation of Anolis lizards in the Greater Antilles; in both cases, relative hindlimb length and perch diameter are strongly correlated. This differentiation could have resulted from genetic change or environmentally-driven phenotypic plasticity. Laboratory studies on A. sagrei from a population in Florida indicate that hindlimb length exhibits adaptive phenotypic plasticity. Further studies are required to determine if the observed differences among the experimental populations are the result of such plasticity. Regardless of whether the differences result from plasticity, genetic change, or both, the observation that anole populations differentiate rapidly and adaptively when exposed to novel environmental conditions has important implications for understanding the adaptive radiation of Caribbean anoles.

The relationship between sexual size dimorphism and habitat use in Greater Antillean Anolis lizards.

Sexual size dimorphism (SSD) is the evolutionary result of selection operating differently on the body sizes of males and females. Anolis lizard species of the Greater Antilles have been classified into ecomorph classes, largely on the basis of their structural habitat (perch height and diameter). We show that the major ecomorph classes differ in degree of SSD. At least two SSD classes are supported: high SSD (trunk-crown, trunk-ground) and low SSD (trunk, crown-giant, grass-bush, twig). Differences cannot be attributed to an allometric increase of SSD with body size or to a phylogenetic effect. A third explanation, that selective pressures on male and/or female body size vary among habitat types, is examined by evaluating expectations from the major relevant kinds of selective pressures. Although no one kind of selective pressure produces expectations consistent with all of the information, competition with respect to structural habitat and sexual selection pressures are more likely possibilities than competition with respect to prey size or optimal feeding pressures. The existence of habitat-specific sexual dimorphism suggests that adaptation of Anolis species to their environment is more complex than previously appreciated.

Effect of predators and area on invasion: an experiment with island spiders.

To assay the effect of area and predators on invasion success, spiders were introduced onto islands that were large, with lizard predators; large, without lizard predators; or small, without lizard predators. Short-term survival was greater on islands without than with predators; area had no effect. Spiders initially increased substantially on both groups of islands without lizards, but after 5 years they nearly died off on small islands while persisting on most large islands; populations in the presence of predators never increased above initial sizes. Results show how predators as well as area are important in determining invasion success.

Lizards reduce food consumption by spiders: mechanisms and consequences.

To determine the effect of lizards on webspider populations, we conducted a long-term field experiment in the Bahamas. Numbers of spider individuals were about 3 times higher in lizard-removal enclosures than in control enclosures with natural densities of lizards. Dietary analyses showed that lizards ate spiders and that lizard and spider diets overlapped substantially. Lizards reduced biomass of prey consumed by spiders; details indicated that they reduced biomass of large (> 4 mm) prey consumed by spiders more than biomass of small (≤4 mm) prey. Similarly, lizards reduced biomass of large aerial arthropods caught in sticky traps but not biomass of small aerial arthropods. We found no evidence that the lizard effect on prey consumption by spiders was caused by a spatial shift from areas with high aerial arthropod abundance to areas with low aerial arthropod abundance. Lizards reduced adult female cephalothorax width and fecundity of spiders. In a separate experiment, food-supplemented spiders were more fecund than control spiders. This study indicates that the interaction between lizards and spiders includes both predation and competition for food.

Leaf pubescence in buttonwood: Community variation in a putative defense against defoliation.

Plants have a variety of putative defenses against defoliation by herbivores, among which are pubescent leaves. Buttonwood (Conocarpus erectus), a Caribbean tree, shows considerable between-individual variation in this trait, and pubescent leaves have less herbivore damage. Surveying 97 island communities, I documented three patterns expected were pubescent individuals more frequent where herbivory is great. (i) Larger islands have a higher percentage of pubescence (larger islands have more herbivores). (ii) Islands nearer to a mainland have a higher percentage of pubescence (nearer islands receive more herbivore immigrants). (iii) Islands having an extremely abundant predator on foliage arthropods, arboreal lizards, have a smaller percentage of pubescence than no-lizard islands. The third effect, though statistically significant, is weak relative to the direct effects of lizards on one category of their prey (spiders) measured in the same system.

Effect of lizards on spider populations: manipulative reconstruction of a natural experiment.

Which species affect one another, how intensely, and the mechanisms of those effects are crucial data for understanding how ecological communities work. Tropical islands without lizards, the major top predators, have about ten times as dense web spider populations as those with lizards; processes responsible for this effect were experimentally simulated by removing lizards from randomly selected mainland plots. Spider densities in removal plots averaged 2.5 times as high as controls. Spider survival, prey abundance, and prey consumption were all negatively affected by lizards. Contrary to most studies, predator removal caused an increase in the number of spider species.

The geographical distribution of rarity.

This paper asks the question: are most species that are censused as rare in particular localities rare throughout most of their geographic ranges, or are they common in substantial portions of their ranges elsewhere? The first alternative is labeled suffusive rarity and the second diffusive rarity. To answer this and similar questions, rarity can be measured as the fraction of censuses from some locality (e.g., a quadrat) in which a species occurs (occurrence rarity), or the relative or absolute abundance of the species averaged over all censuses from some locality (abundance rarity). The question was analyzed for occurrence-rarity data from Australian terrestrial birds distributed over 1° (104-km2) quadrats. The great majority of species that are rare in a particular quadrat are not rare and are often common in a substantial number of other quadrats, i.e., these avian species are much closer to the diffusive than suffusive portion of the rarity continuum. The data also show that 1) the distribution of sizes of geographic ranges, whether breeding or total, is highly skewed, appearing exponential to more concave; 2) species are much rarer in their nonbreeding than breeding ranges; 3) more widespread species, whether breeding or total ranges are considered, tend to occur more rarely in a slightly but significantly greater fraction of their ranges; and 4) hawks and owls, typified by high abundance rarity, show occurrence rarity in a greater fraction of their ranges than the average nonraptorial species. Although continental birds may be especially predilected toward diffusive rarity, the present analysis points to identification of centers of abundance as major ways of preserving those species contributing most to recorded instances of rarity. Similar analyses with other kinds of organisms would be most welcome.

Spider populations: extraordinarily high densities on islands without top predators.

Distributions and densities of orb spiders on small islands are extremely variable. Species occurrences are far more irregular for spiders than vertebrates on the same islands. Much variation in spider density is explainable by distance from the presumed source of colonists and presence or absence of vertebrate predators. As has been predicted for passive dispersers, densities decline exponentially with distance. For a given distance, spider densities are about ten times greater on islands without vertebrate predators than on those with such predators.

The compression hypothesis and temporal resource partitioning.

Contingency models of feeding compare the energy per unit time gained from utilizing a resource unit of a particular kind (food types, habitat patches, time periods) against that energy/time expected if the unit is skipped. Optimally, an animal should reject the particular unit if and only if the former energy/time is less than the latter. Consequently, food or habitat types should be excluded if the prospect of finding and consuming better types is sufficiently high. In contrast, feeding periods should be skipped only if it is less costly to wait than to feed.In situations of high food abundance, contingency models imply that animals should be maximally specialized with respect to food or habitat type, but maximally generalized with respect to time period. As food decreases in abundance, food and habitat types should be added to the diet or itinerary, but time periods should be omitted from feeding activity. In contrast, animals with fixed caloric intake should broaden diet, habitat, and feeding times as abundance decreases.According to contingency models, competitors cannot cause item kinds to be dropped from the diet, but because they can affect the values of patches once found, can cause habitat kinds to be dropped from the itinerary. Competitors also reduct the value of feeding during particular time periods, but ordinarily fairly severe depletion must occur before it is optimal to feed no longer in a period frequented by competitors.These arguments imply that temporal resource partitioning on a diel basis should be relatively rare. In fact, specialization according to feeding period should differentially occur in animals of limited abilities to use or process available food, whereas that need not be the case for food-type or habitat specialization. It should also occur in animals sensitive to and found in variable climates.

Resource partitioning in ecological communities.

To understand resource partitioning, essentially a community phenomenon, we require a holistic theory that draws upon models at the individual and population level. Yet some investigators are still content mainly to document differences between species, a procedure of only limited interest. Therefore, it may be useful to conclude with a list of questions appropriate for studies of resource partitioning, questions this article has related to the theory in a preliminary way. 1) What is the mechanism of competition? What is the relative importance of predation? Are differences likely to be caused by pressures toward reproductive isolation? 2) Are niches (utilizations) regularly spaced along a single dimension? 3) How many dimensions are important, and is there a tendency for more dimensions to be added as species number increases? 4) Is dimensional separation complementary? 5) Which dimensions are utilized, how do they rank in importance, and why? How do particular dimensions change in rank as species nuimber increases? 6) What is the relation of dimensional separation to difference in phenotypic indicators? To what extent does the functional relation of phenotype to resource characteristics constrain partitioning? 7) What is the distance between mean position of niches, what is the niche standard deviation, and what is the ratio of the two? What is the niche shape?

The Ecological Significance of Sexual Dimorphism in Size in the Lizard Anolis conspersus.

Adult males of Anolis conspersus capture prey of significantly larger size and occupy perches of significantly greater diameter and height than do adult females; similarly, these three dimensions of the niche are significantly larger for adult females than for juveniles. Adult males on the average eat a smaller number of prey, and the range in size of prey is larger. The relationship between the average length of the prey and that of the predator is linear when the predator size is above 36 millimeters, but becomes asymptotic when it is below that value. Subadult males as long as adult females eat significantly larger food than do the latter, but only in the larger lizards is this correlated with a relatively larger head. Anolis conspersus selects prey from a wide range of taxa and shows no obvious intraspecific specialization not connected to differences in microhabitat and prey size. The efficiency of this system for solitary species is pointed out.