Yaron Ziv

Interference competition and temporal and habitat partitioning in two gerbil species

We have tested two hypotheses which may explain two different patterns which underlie coexistence in two species of desert gerbils (Gerbillus allenbyi and G. pyramidum). The two patterns are temporal partitioning of foraging activity and shared preference habitat selection. When sympatric, G. pyramidum uses the early part of the night most heavily while G. allenbyi is most active in the late part of the night. Although both species prefer the same habitat type (semistabilized sand dune), in the presence of G, pyramidum, G. allenbyi utilizes mainly the secondary habitat type (stabilized sand dune)

Inferring the history of speciation in house mice from autosomal, X-linked, Y-linked and mitochondrial genes

Patterns of genetic differentiation among taxa at early stages of divergence provide an opportunity to make inferences about the history of speciation. Here, we conduct a survey of DNA‐sequence polymorphism and divergence at loci on the autosomes, X chromosome, Y chromosome and mitochondrial DNA in samples of Mus domesticus, M. musculus and M. castaneus. We analyzed our data under a divergence with gene flow model and estimate that the effective population size of M. castaneus is 200 000–400 000, of M. domesticus is 100 000–200 000 and of M. musculus is 60 000–120 000. These data also suggest that these species started to diverge approximately 500 000 years ago. Consistent with this recent divergence, we observed considerable variation in the genealogical patterns among loci. For some loci, all alleles within each species formed a monophyletic group, while at other loci, species were intermingled on the phylogeny of alleles. This intermingling probably reflects both incomplete lineage sorting and gene flow after divergence. Likelihood ratio tests rejected a strict allopatric model with no gene flow in comparisons between each pair of species. Gene flow was asymmetric: no gene flow was detected into M. domesticus, while significant gene flow was detected into both M. castaneus and M. musculus. Finally, most of the gene flow occurred at autosomal loci, resulting in a significantly higher ratio of fixed differences to polymorphisms at the X and Y chromosomes relative to autosomes in some comparisons, or just the X chromosome in others, emphasizing the important role of the sex chromosomes in general and the X chromosome in particular in speciation.

REGIONAL VARIATION IN THE HIERARCHICAL PARTITIONING OF DIVERSITY IN CORAL‐DWELLING FISHES

The size of the regional species pool may influence local patterns of diversity. However, it is unclear whether certain spatial scales are less sensitive to regional influences than others. Additive partitioning was used to separate coral-dwelling fish diversity to its alpha and beta components, at multiple scales, in several regions across the Indo-Pacific. We then examined how the relative contribution of these components changes with increased regional diversity. By employing specific random-placement null models, we overcome methodological problems with local-regional regressions. We show that, although alpha and beta diversities within each region are consistently different from random-placement null models, the increase in beta diversities among regions was similar to that predicted once heterogeneity in coral habitat was accounted for. In contrast, alpha diversity within single coral heads was limited and increased less than predicted by the null models. This was correlated with increased intraspecific aggregation in more diverse regions and is consistent with ecological limitations on the number of coexisting species at the local scale. These results suggest that, apart from very small spatial scales, variation in the partitioning of fish diversity along regional species richness gradients is driven overwhelmingly by the corresponding gradients in coral assemblage structure.

Habitat fragmentation may not matter to species diversity

Conservation biologists worry that fragmenting a bloc of natural habitat might reduce its species diversity. However, they also recognize the difficulty and importance of isolating the effect of fragmentation from that of simple loss of area. Using two different methods (species–area curve and Fishers α index of diversity) to analyse the species diversities of plants, tenebrionid beetles and carabid beetles in a highly fragmented Mediterranean scrub landscape, we decoupled the effect of degree of fragmentation from that of area loss. In this system, fragmentation by itself seems not to have influenced the number of species. Our results, obtained at the scale of hectares, agree with similar results at island and continent scales.

Costs of two non-mutualistic species in a yucca/yucca moth mutualism

Mutualisms often involve significant costs for participants. Costs are inflicted by mutualists themselves, as well as by associated, non-mutualistic species. These costs are rarely quantified, however, particularly the ones extrinsic to the pairwise interaction. We compare costs inflicted by an obligate mutualist pollinator and two common exploiters of an Arizona yucca over a 2-year period. The magnitude of seed damage from seed and fruit-feeding beetle larvae (Carpophilus longus, Nitidulidae) was similar to damage from the seed-eating larvae of Yucca schottiis pollinator moth Tegeticula yuccasella (Prodoxidae), averaging about 15 seeds destroyed per fruit in each case. The two seed predators usually fed within the same fruits, although rarely side by side. In contrast, the presence of fruit-galling moth larvae (Prodoxusy-inversus, Prodoxidae) appeared to benefit the yucca: individual Tegeticula destroyed only half as many seeds in galled fruits as they did in ungalled fruits. We discuss three general implications of these results. Firstly, the costs of non-mutualists to the two mutualistic partners are not necessarily parallel. Secondly, measurable costs of non-mutualists do not necessarily translate into an impact on the success of the mutualism itself, because they may be incurred after mutualistic activities take place. Finally, the costs of mutualists to each other can differ substantially depending on the presence or absence of non-mutualistic species.

Infertile seeds ofYucca schottii: A beneficial role for the plant in the yucca-yucca moth mutualism?

SummaryThe yucca-yucca moth interaction is a classic case of obligate mutualism. Female moths pollinate and oviposit in the gynoecium of the flower; however, maturing larvae eat a fraction of the developing seeds. We studied within-fruit distributions of four seed types (fertile, infertile, eaten and uneaten seeds) in order to evaluate costs and benefits in aYucca schottii population in southeastern Arizona. We focused on how the spatial arrangement of seeds affected larval behaviour and, hence, the costs of the mutualism to the yucca. Infertile seeds were distributed throughout both infested and uninfested locules. Additionally, moth larvae feeding in a single locule preferred fertile seeds and even avoided infertile seeds and left the fruit significantly more often when they encountered infertile seeds. We suggest that, regardless of the cause of infertile seeds, they function as blocking units within seed locules and therefore reduce seed predation by moth larvae. We also suggest that, together with certain other fruit traits, the presence of infertile seeds promotes the evolutionary stability of this pollination mutualism.

Decoupling fragmentation from habitat loss for spiders in patchy agricultural landscapes.

Habitat loss reduces species diversity, but the effect of habitat fragmentation on number of species is less clear because fragmentation generally accompanies loss of habitat. We compared four methods that aim to decouple the effects of fragmentation from the effects of habitat loss. Two methods are based on species-area relations, one on Fishers alpha index of diversity, and one on plots of cumulative number of species detected against cumulative area sampled. We used these methods to analyze the species diversity of spiders in 2, 3.2 × 4 km agricultural landscapes in Southern Judea Lowlands, Israel. Spider diversity increased as fragmentation increased with all four methods, probably not because of the additive within-patch processes, such as edge effect and heterogeneity. The positive relation between fragmentation and species diversity might reflect that most species can disperse through the fields during the wheat-growing season. We suggest that if a given area was designated for the conservation of spiders in Southern Judea Lowlands, Israel, a set of several small patches may maximize species diversity over time.

Giving-up densities of foraging gerbils: the effect of interspecific competition on patch use

We studied the effect of a dominant species, Gerbillus pyramidum (Egyptian sand gerbil), on the patch use of its subordinate competitor, G. andersoni allenbyi (Allenbys gerbil), to better understand interspecific competition between the two species. We used manipulated resource patches (seed trays) covered with cages with two adjustable species-specific gates (either opened or closed to the bigger-dominant species, but always opened to the subordinate one). We recorded species tracks around and on the seed trays and giving-up densities (GUDs) of seeds in the trays after each night of foraging. G. a. allenbyi depleted seed patches to a lower level whenever G. pyramidum was given the opportunity to forage on the seed trays (i.e., present on the grid). This result held regardless of whether G. pyramidum was actually present at a particular station. We suggest that competition from G. pyramidum occurs both directly by interference, in which G. a. allenbyi is forced to be active in the late part of the night, and indirectly by exploitation via resource depletion by G. pyramidum in the early part of the night. The results suggest that interspecific competition from G. pyramidum reduces seed availability and the richness of the environment for G. a. allenbyi enough to affect the marginal value of energy for G. a. allenbyi individuals and cause them to experience lower costs of predation and manifest lower GUDs.

Determinants of the steep species–area relationship of coral reef fishes

The increase in species richness with area is known as the species–area relationship (SPAR). Although several mutually non-exclusive processes may produce the SPAR, the null, often ignored, hypothesis states that a SPAR can be generated by random placement alone. The log–log-transformed SPAR of coral reef fishes on small patch-reefs revealed a steep slope of 0.55. However, this slope was dependent on the cumulative area of the reef examined and was therefore affected by random placement. After statistically removing the contribution of random placement from the SPAR, the slope was estimated to be 0.21. This is consistent with estimates from other, mostly terrestrial, systems. Furthermore, a randomization procedure, where the probability of fishes to reach a patch was proportional to reef area, showed that the field measured SPAR did not differ from random placement. In addition, fish assemblages on species poor reefs did not form subsets of species rich reefs (i.e., no nestedness) beyond that expected from random placement. Steep log–log-transformed SPARs can be formed by random placement alone, indicating that caution should be used when assigning an ecological meaning to SPARs generated from small spatial scales.

ON THE SCALING OF HABITAT SPECIFICITY WITH BODY SIZE

Larger species tend to occupy more habitats, but a theoretical framework for the pattern is lacking. I modified the continuous-time logistic equation of population growth in two ways to allow for such a habitat-based theoretical framework. First, I separated birth rate from death rate. Second, I included two new terms in the equation: (1) an explicit spatial variable for habitat quality that reflects the match between a habitat and a population (species–habitat match), and (2) a demand/supply function that depends on the ratio between the energy used by all populations occurring in a habitat, and energy available in that habitat. Energy was used as a common currency to overcome differences between species of different body sizes as well as to overcome differences caused by disproportional intra- and interspecific effects. Allometric relations were used to characterize parameter values that correlate with body size, such as metabolic rate, birth rate, and death rate. The analytical solution of the equation for carrying capacity shows that, for a population to have a positive carrying capacity, its ratio of death rate to birth rate should be less than its match to the habitat it occupies. Literature-based body-size-dependent birth and death rates of Eutherian mammals show that the death-rate:birth-rate ratio decreases with body size. Combining the analytical solution and the death-rate:birth-rate ratio reveals that habitat generality should positively scale with body size. I used this model to simulate simple spatially explicit landscapes having diverse habitats and combinations of species of various body sizes. Using realistic parameters, the model generates results that are consistent with field observations. Thus, one can focus on specific processes to explore macroecological questions.