The role of phylogeny in desert rodent community assembly

Recent advances in molecular genetics and phylogenetic reconstruction have the potential to transform ecology by providing new insights into the historical evolution of ecological communities. This study by Stevens and collaborators complements decades of previous research on desert rodents, by combining data from a field study and a phylogenetic tree for Mojave Desert rodents to address patterns and processes of community assembly. The number of coexisting rodent species is positively correlated, and the average phylogenetic distance among these species is negatively correlated with perennial plant species richness. As rodent species diversity increases along a gradient of increasing environmental heterogeneity, communities are composed of increasingly related species: there is a consistent pattern of phylogenetic structure from over-dispersed through random to clumped. I discuss this pattern in the light of complementary results of previous studies. This paper is noteworthy for calling attention to still unanswered questions about how the historical events of speciation, colonization, extinction, and trait evolution and their relationship to past climates and vegetation have given rise to current patterns of community organization.     

Structure in a desert rodent community: use of space around Dipodomys spectabilis mounds

Summary In an earlier paper (Bowers et al. 1987) we reported patterns of microhabitat use by desert rodents among 0.25-ha plots where seeds were added or certain rodent species removed. We used the results to make inferences about the spatial organization of the whole rodent community. Here we change our focus to test for spatial usage patterns at a smaller (within-plot) scale. Specifically, we examine to what extent spatial use varies with proximity to mounded burrows of the large kangaroo rat, Dipodomys spectabilis. Capture frequency of five of nine rodent species was correlated with distance from D. spectabilis mounds, while six species showed correlations with vegetative cover that also increased with distance from mounds. In general, sites nearer mounds were underutilized, and sites further away overutilized by the rodent community as a whole. Logistic regression analyses showed for six species that vegetative cover and particularly, distance to nearest mound accounted for more of the variation in whether a station captured a species than did plot-to-plot (i.e., treatment) effects. Similar analyses using two plots where D. spectabilis was removed (where vacant mounds persited for years) suggested that the selection of microhabitats with contrasting vegetation structure was more pronounced in the absence than in the presence of D. spectabilis, and that in most cases distance from mounds was important only if mounds were occupied. Spatial structure in this community appears to revolve around the occupation of space by dominant individuals that partially modify/obscure large scale patterns involving the selection of particular structural microhabitats.     

Spatial organization of a desert rodent community: food addition and species removal

Summary From 1977 through 1983 we conducted experiments on a desert rodent community where supplemental seeds were added or certain rodent species and ants were removed from 0.25-ha fenced plots in a Chihuahuan Desert site in southeastern Arizona, USA. In this paper we examine the patterns of microhabitat use relative to vegetative cover by 11 rodent species. The results show that: i) removal of the largest seed-eating species, Dipodomys spectabilis, produced the most pervasive and dramatic shifts in microhabitat use by the remaining rodent species; ii) adding seeds or removing ants had little effect on the spatial use of microhabitats by rodents in this community; and iii) non-granivores were just as likely as granivores to shift microhabitat use when other granivores were removed. We believe these results indicate that both food and foraging microsites are limited but the relegation of subdominant species to lesspreferred microhabitats by the large Dipodomys spectabilis is the major factor underlying the spatial organization of this community. Results also demonstrate that strong interactions among species increase the probability that pathways of indirect interactions through intermediary species are important; these complex linkages may include species that overlap little in food preferences.     

Response of lizard community structure to desert grassland restoration mediated by a keystone rodent

Many grasslands in the Chihuahuan Desert have transformed to shrublands dominated by creosotebush (Larrea tridentata). Grassland restoration efforts have been directed at controlling creosotebush by applying herbicide over large spatial scales. However, we have a limited understanding of how landscape-scale restoration affects biodiversity. We examined whether restoration treatments in southern New Mexico, USA have influenced the community structure of lizards, which are sensitive to shrub encroachment. We compared lizard community structure on 21 areas treated with herbicide from 7 to 29 years ago with paired untreated areas that were dominated by shrubs and matched by geomorphology, soils, and elevation. To examine mechanisms underlying responses to restoration, we tested whether the abundance of a grassland specialist, Aspidoscelis uniparens, depended on time since treatment, treatment area and isolation, and local habitat quality. Because lizards use rodent burrows as habitat, we tested whether community structure and A. uniparens abundance depended on the abundance of the keystone rodent, Dipodomys spectabilis. Treated areas had reduced shrub cover and increased grass cover compared to untreated areas. Lizard community composition differed strongly between areas, with four species responding to treatments. Divergence in community composition between treated–untreated pairs was greatest for old treatments (≥22 years), and community composition was influenced by D. spectabilis. In particular, the abundance of A. uniparens was greatest on old treatments with a high density of D. spectabilis. Overall, our results demonstrate lizard community structure responds to grassland restoration efforts, and keystone species can shape restoration outcomes. Reestablishment of keystone species may be a critical constraint on the recovery of animal biodiversity after habitat restoration.     

Ectoparasites and age-dependent survival in a desert rodent

Host age is one of the key factors in host–parasite relationships as it possibly affects infestation levels, parasite-induced mortality of a host, and parasite distribution among host individuals. We tested two alternative hypotheses about infestation pattern and survival under parasitism in relation to host age. The first hypothesis assumes that parasites are recruited faster than they die and, thus, suggests that adult hosts will show higher infestation levels than juveniles because the former have more time to accumulate parasites. The second hypothesis assumes that parasites die faster than they are recruited and, thus, suggests that adults will show lower infestation levels because of acquired immune response and/or the mortality of heavily infested juveniles and, thus, selection for less infested adults. As the negative effects of parasites on host are often intensity-dependent, we expected that the age-related differences in infestation may be translated to lower or higher survival under parasitism of adults, in the cases of the first and the second hypotheses, respectively. We manipulated ectoparasite numbers using insecticide and assessed the infestation pattern in adult and juvenile gerbils (Gerbillus andersoni) in the Negev Desert. We found only a partial support for age-dependent parasitism. No age-related differences in infestation and distribution among host individuals were found after adjusting the ectoparasite numbers to the host’s surface area. However, age-related differences in survival under parasitism were revealed. The survival probability of parasitized juveniles decreased in about 48% compared to unparasitized hosts while the survival probability of adults was not affected by ectoparasites. Our results suggest that the effect of host age on host–parasite dynamics may not explicitly be determined by age-dependent differences in ectoparasite recruitment or mortality processes but may also be affected by other host-related and parasite-related traits.     

Spatial structure and niche patterns of a rodent community in the south Bukhara desert (Middle Asia)

The distribution of niches in resource space and the niche patterns of a 13 species community of Middle Asian desert rodents were studied by use of discriminant function analysis Nineteen quantitative parameters of the environment (soil structure and vegetation), measured at 600 sample plots within twenty four one ha grids, were considered The first three canonical axes of resource space account for 83% of the variance, the first axis represents a general landscape gradient from sand to clay soils, the second axis reflects a gradient of increasing productivity, and the third axis reflects a gradient of increasing protectability of the environment The distribution of niches in resource space is not even, there are two distinct spatial guilds consisting of psammophilous and sclerophilous species There is a negative correlation between niche position (distance from species centroid to the center of the resource space) and maximal population biomass At the same time there are no correlations between the niche breadth and the niche position due to the absence of species with really broad niches Rodent biomass increases along the productivity axis and reaches a peak in the middle part of the substrate axis Rodent species diversity increases along the substrate axis from sand to clay soils Changes of species diversity along productivity axis have humpshaped patterns Maximum species diversity was recorded at low level of productivity on the sandy soils and shifted to intermediate levels of productivity on sandy-loam and clay soils     

Tadpole competition in a desert anuran community

Summary Five interspecific competition experiments performed in artificial ponds under temporary pond-like conditions established that temporary and permanent pond tadpoles interact. Temporary pond species typically lowered survivorship and reduced progress towards metamorphosis of permanent pond tadpoles. These results suggest that competitive interactions may partially determine the local distribution of tadpoles in the Chihuahuan Desert by preventing some permanent pond anurans from using temporary ponds. Predation by Scaphiopus multiplicatus tadpoles (a temporary pond species) may also prevent permanent pond tadpoles from using temporary ponds.     

Developmental and acclimatory contributions to water loss in a desert rodent: investigating the time course of adaptive change

Understanding the evolution of physiological traits requires considering three nonexclusive mechanisms that underlie phenotypes and cause their change over different time scales: acclimation, developmental plasticity, and natural selection for genetically fixed traits. Physiological adjustments to changes in the desiccating potential of the environment were investigated with one subspecies of common desert rodent, Dipodomys merriami merriami (Merriam's kangaroo rat). We raised young whose parents originated from environments that differ in both temperature and humidity. These young were raised under either desiccating or water-abundant conditions, and their water loss was measured at a series of temperatures to determine the effect developmental conditions have on resistance to desiccation. We then determined the contribution of acclimation to desiccation resistance by keeping the differentially raised young in conditions opposite to those during their development and again measuring water loss. We found that developmental plasticity and acclimation can completely account for the existing intraspecific variability in desiccation resistance under certain conditions. In fact, developmental and acclimatory changes can equal genetically based differences of the populations. This phenotypic plasticity can operate relatively quickly and therefore may attenuate the actions of natural selection. Understanding the extent and nature of such flexibility is critical to our understanding intraspecific variability and the consequences of changing climate.     

Clump spacing in a desert dwarf shrub community

A Monte Carlo method based on Ripley's K function-a cumulative function related to the number of plants encountered at different distances from other plants-is used to test the null hypothesis of random distribution of shrub clumps in a desert dwarf shrub community in Namaqualand, South Africa, where Psilocaulon arenosum is the dominant shrub. The method takes into account the apparent regularity of pattern caused by the finite size (up to 2 m diameter) of the clumps. It is shown that the clump centres are significantly aggregated (compared to random expectation) at distances on the order of 1 m. Such aggregation is expected, as a simple result of regeneration near to seed sources, if the time between catastrophic droughts is short in relation to the time required for development of a non-aggregated or regular pattern determined by moisture competition. No significant regulatiry was detected at distances of 3 m or less. One subplot showed regularity above 3 m, but this pattern was not shown by the other subplot and may not be a competition effect. These results support a hypothesis of aggregation caused by regeneration pattern decaying slowly toward randomness as larger individuals compete.We thank C. J. F. ter Braak for suggesting the method of spatial analysis; 1. Noy-Meir, C. J. F. ter Braak and two unnamed reviewers for helpful comments; the Netherlands Science Research Council (ZWO) for financial support to I. C. Prentice (grant B84-218); and Prof. D. von Willert for material support to M. J. A. Werger.     

The metabolism of phenol in a desert rodent Notomys alexis.

1. Phenol is rapidly metabolized in the desert rodent Notomys alexis; sulphate and glucuronide conjugates being formed.2. The circulating half-life of phenolic metabolites is similar in N. alexis to laboratory mice. This contrasts with comparable data for salicylate turnover.3. After injection, the tissue distribution of phenolic metabolites is highest in liver, lung and kidney.4. In this species, phenol is rapidly conjugated to soluble form and a low rate of water turnover does not correlate with differences in phenol metabolism from those generally observed in mammals.     

Trade-offs between seed dispersal and dormancy in an amphi-basicarpic cold desert annual

Background and Aims

Several studies have demonstrated trade-offs between depth of seed dormancy and dispersal ability for diaspore-dimorphic species. However, relatively little is known about trade-offs between these two life history traits for a species that produces more than two diaspore morphs. The aim of this study was to investigate the relationship between seed dormancy and dispersal in Ceratocarpus arenarius, an amphi-basicarpic cold desert annual that produces a continuum of dispersal unit morphs.

Methods

A comparison was made of dispersal and dormancy breaking/germination responses of dispersal units from ground level (a), the middle of the plant canopy (c) and the top of the plant canopy (f). Various features of the morphology and mass of dispersal units and fruits (utricles) were measured. The role of bracteoles in diaspore dispersal by wind, settlement onto the soil surface and dormancy/germination was determined by comparing responses of intact dispersal units and fruits. Movement of dispersal units by wind and animals, seed after-ripening, germination phenology and the presence of water-soluble germination inhibitors in bracteoles were tested using standard procedures.

Key Results

Dispersal units a, c and f differed in morphology and mass; in the majority of cases, extremes were exhibited by a and f, with c being intermediate. Overall, relative dispersal ability was f > c > a, whereas relative intensity of dormancy was a > c > f. Bracteoles increased dispersal distance by wind, enhanced settlement of diaspores onto the soil surface and mechanically inhibited germination.

Conclusions

The results provide evidence for a model in which there is a continuous inverse-linear relationship between diaspore dispersal ability and depth of dormancy. Thus, dispersal unit heteromorphism of C. arenarius results in a continuum, from no dispersal ability/high dormancy (dispersal unit a) to high dispersal ability/low dormancy (unit f), which may be a bet-hedging strategy in the cold desert environment.     

Ecological organization of a Chihuahuan desert lizard community

Summary Niche relationships among 11 diurnal lizard species were studied for 3 years. The community appeared to be subdivided spatially into different subsets of only four to six species. Coexisting species of each subgroup were separated further by differential utilization of microhabitats. In late spring and early summer, when food shortage is likely, dietary overlaps between syntopic species were generally low but increased broadly after the rains, when food became plentiful. Because of the high similarity in prey-size utilization among large and medium species, effects of prey selection according to size were generally slight except for very different-sized species. Overlap in the three-dimensional resource space was high between Cnemidophorus scalaris and C. tigris, but close observation of the ecology of the two populations showed many small differences between them. We suggest that the very diversified ecological opportunism of C. scalaris allows it to coexist with C. tigris in a very heterogeneous ecosystem. Several hypotheses are discussed to explain the ecological determinism of resource partitioning within such a community.     

Spatial patterns of mortality in a Colorado desert plant community

Spatial pattern and patterns of mortality were analyzed from a five year re-census of over 7000 mapped perennial shrubs within a one hectare study plot in Joshua Tree National Park. A five year interval revealed evidence for negative intraspecific interactions which was not evident after a one year evaluation of the same plants within the study area. Juvenile mortality was a positive function of conspecific density for Ambrosia dumosa, Eriogonum fasciculatum and Sphaeralcea ambigua. In addition, juvenile E. fasciculatum that died occurred closer to conspecific adults than would be expected by chance. The two censuses are similar in that uniform patterns of distribution among conspecifics are rare, particularly for non-succulent species.These results emphasize the value of mapped study sites in allowing dynamic inferences from otherwise static analyses. Herbivory and nurse effects are implicated as potential factors influencing the dispersion of perennial plants within deserts.     

Trade-offs in skillacquisition and time allocation among juvenile chacma baboons

We hypothesize that juvenile baboons are less efficient foragers than adult baboons owing to their small size, lower level of knowledge and skill, and/or lesser ability to maintain access to resources. We predict that as resources are more difficult to extract, juvenile baboons will demonstrate lower efficiency than adults will because of their lower levels of experience. In addition, we hypothesize that juvenile baboons will be more likely to allocate foraging time to easier-to-extract resources owing to their greater efficiency in acquiring those resources. We use feeding efficiency and time allocation data collected on a wild, free-ranging, non-provisioned population of chacma baboons (Papio hamadryas ursinus) in the Moremi Wildlife Reserve, Okavango Delta, Botswana to test these hypotheses. The major findings of this study are: 1. Juvenile baboons are significantly less efficient foragers than adult baboons primarily for difficult-to-extract resources. We propose that this age-dependent variation in efficiency is due to differences in memory and other cognitive functions related to locating food resources, as is indicated by the greater amount of time juvenile baboons spend searching for food. There is no evidence that smaller body size or competitive disruption influences the differences in return rates found between adult and juvenile baboons in this study. 2. An individual baboon’s feeding efficiency for a given resource can be used to predict the duration of its foraging bouts for that resource. These results contribute both to our understanding of the ontogeny of behavioral development in nonhuman primates, especially regarding foraging ability, and to current debate within the field of human behavioral ecology regarding the evolution of the juvenile period in primates and humans. Sara E. Johnson is Assistant Professor of Anthropology at California State University, Fullerton. She received her Ph.D. in Anthropology (Human Evolutionary Ecology) from the University of New Mexico in 2001. She uses behavioral ecology and life history theory to address her research interests in the evolution of primate and human growth; ecological variation and phenotypic plasticity in growth and development; ecological variation in life course trajectories, including fertility, health, morbidity, and mortality differentials; food acquisition and production related to nutrition; societal transofmration and roles of the elderly among indigenous peoples; and women’s reproductive and productive roles in both traditional and nontraditional societies. For the past decade she has conducted research on these issues in several different populations, including chacma baboons in the Okavango Delta of Botswana, two multiethnic communities of forager/agropastoralists in the Okavango Delta of Botswana, and among New Mexican men. John Bock is Associate Professor of Anthropology at California State University at Fullerton and is Associate Editor of Human Nature. He received a Ph.D. in Anthropology (Human Evolutionary EcologY) from the University of New Mexico in 1995, and from 1995 to 1998 was an Andrew W. Mellon Foundation postdoctoral fellow in demography and epidemiology at the National Centre for Epidemiology and Population Health at Australian National University. His recent research has focused on applying life history theory to understanding the evolution of the primate and human juvenile period. Bock has been conducting research among the Okavango Delta peoples of Botswana since 1992, and his current research there is an examination of child development and family demography in relation to socioecology and the HIV/AIDS epidemic. Other research is focused on health disparties among minorities and indigenous peoples in Botswana and the United States related to differential access to health care.     

Species and plant community distribution in a Mojave-Great Basin desert transition

Multivariate analyses were used to describe the vegetation characteristics of a transition from lowelevation Mojave desert to higher-elevation Great Basin desert. Vegetation data used were from Plutonium Valley in the Nevada Test Site. Data from forty nine releves were analyzed with two classifications (two-day indicator analysis or TWINSPAN and unweighted paried group cluster analysis or CLUSTER). Three ordinations, reciprocal averaging (RA), detrended reciprocal averaging (DCA) and non-metric multidimensional scaling (MNDS), were also used. A rotational correlation analysis was used to determine the vector direction of environmental gradients that correlate best with ordination results. Only token correspondence was found between multivariate classes generated by TWINSPAN and CLUSTER, and seven classes (plant communities) identified from field reconnaissance. The latter seven communities were based on differences in dominant species. Distribution of the vegetation was related more to beta diversity than alpha diversity. Individual species were much less diagnostic than the amount of plant cover, groups or guilds of species or differences in elevation and steepness of slope. Because of the high beta diversity the NMDS ordination gave results with the greatest ease of interpretation.