Susan L. Durham

On the use of log‐transformation vs. nonlinear regression for analyzing biological power laws

Power-law relationships are among the most well-studied functional relationships in biology. Recently the common practice of fitting power laws using linear regression (LR) on log-transformed data has been criticized, calling into question the conclusions of hundreds of studies. It has been suggested that nonlinear regression (NLR) is preferable, but no rigorous comparison of these two methods has been conducted. Using Monte Carlo simulations, we demonstrate that the error distribution determines which method performs better, with NLR better characterizing data with additive, homoscedastic, normal error and LR better characterizing data with multiplicative, heteroscedastic, lognormal error. Analysis of 471 biological power laws shows that both forms of error occur in nature. While previous analyses based on log-transformation appear to be generally valid, future analyses should choose methods based on a combination of biological plausibility and analysis of the error distribution. We provide detailed guidelines and associated computer code for doing so, including a model averaging approach for cases where the error structure is uncertain.

Species Interactions at the Level of Fine Roots in the Field: Influence of Soil Nutrient Heterogeneity and Plant Size

Interference at the level of fine roots in the field was studied by detailed examination of fine root distribution in small soil patches. To capture roots as they occur in natural three-dimensional soil space, we used a freezing and slicing technique for microscale root mapping. The location of individual roots intersecting a sliced soil core surface was digitized and the identity of shrub and grass roots was established by a chemical technique. Soil patches were created midway between the shrub, Artemisia tridentata, and one of two tussock grasses, Pseudoroegneria spicata or Agropyron desertorum. Some soil patches were enriched with nutrients and others given only deionized water (control); in addition, patches were located between plants of different size combination (large shrubs with small tussock grasses and small shrubs with large tussock grasses). The abundance of shrub and grass roots sharing soil patches and the inter-root distances of individual fine roots were measured. Total average rooting density in patches varied among these different treatment combinations by only a factor of 2, but the proportion of shrub and grass roots in the patches varied sixfold. For the shrub, the species of grass roots sharing the patches had a pronounced influence on shrub root density; shrub roots were more abundant if the patch was shared with Pseudoroegneria roots than if shared with Agropyron roots. The relative size of plants whose roots shared the soil patches also influenced the proportion of shrub and grass roots; larger plants were able to place more roots in the patches than were the smaller plants. In the nutrient-enriched patches, these influences of grass species and size combination were amplified. At the millimeter- to centimeter-scale within patches, shrub and grass roots tended to segregate, i.e., avoid each other, based on nearest-neighbor distances. At this scale, there was no indication that the species-specific interactions were the result of resource competition, since there were no obvious patterns between the proportion of shrub and grass roots of the two species combinations with microsite nutrient concentrations. Other potential mechanisms are discussed. Interference at the fine-root level, and its species-specific character, is likely an influential component of competitive success, but one that is not easily assessed.

Susceptibility of pollen to UV-B radiation: an assay of 34 taxa.

Much of the ultraviolet-B radiation (UV-B) research on plants has concentrated on vegetative plant parts, and only a small fraction has dealt with the reproductive system. The present study analyzed pollen grains of 34 taxa germinated and grown under two levels of UV-B radiation (187 and 460 mW/m2) or no UV-B (control group). Visible radiation at 260 mmol/m/s was present in all treatments. Taxa included those with binucleate and trinucleate pollen types. We detected differences among species. A significant reduction in pollen germination occurred in only five species. Pollen tubes of >50% of the species showed significant reduction in length. Trinucleate pollen types were more likely to exhibit tube length reduction than the binucleate types. Proportionately more monocotyledonous species were sensitive to UV-B treatment than dicotyledonous species, and proportionately more wild species were sensitive than cultivated species and pollen collected from plants growing in the field were somewhat more sensitive than pollen collected from plants grown in the greenhouse. Species in which pollination occurred earlier in the season were more likely to be susceptible to UV-B radiation than those for which anthesis took place later in the season, suggesting a possible adaptation to UV-B radiation.

EFFECTS OF NUTRIENT PATCHES AND ROOT SYSTEMS ON THE CLONAL PLASTICITY OF A RHIZOMATOUS GRASS

Clonal plant foraging has been examined primarily on individual clones exposed to resource-poor and resource-rich environments. We designed an experiment to examine the clonal foraging behavior of the rhizomatous grass Elymus lanceolatus ssp. lanceolatus under the influence of neighboring plant root systems in a heterogeneous nutrient environment. Individual Elymus clones were planted in large bins together with one of three neighboring grass species, Agropyron desertorum, Pseudoroegneria spicata, or Bromus tectorum, which differ in rooting density and growth activity. The position of Elymus clones was manipulated so rhizomes encountered a short-duration nutrient patch and subsequently root systems of the neighboring plants. Unexpectedly, the morphological plasticity of the perennial grass Elymus lanceolatus ssp. lanceolatus was influenced by the presence of the neighboring species much more than by the local nutrient enrichments, although nutrient patches did amplify some of the foraging responses. Elymus rhizomes branched readily and initiated large daughter plants as they encountered the low-density root systems of Pseudoroegneria. When Elymus encountered the fine, dense root systems of the annual Bromus, clonal expansion was initially reduced. Yet, after the short growing season of Bromus, Elymus resumed clonal expansion and produced several daughter plants. Elymus clones were most constrained by the fine, dense root systems of Agropyron desertorum. In this case, a few, long rhizomes avoided the densely rooted soil environment by growing aboveground as stolons crossing over the Agropyron tussocks. Elymus clonal biomass was largest in neighborhoods of Pseudoroegneria, intermediate in neighborhoods with Bromus, and smallest in neighborhoods with Agropyron. The latter were approximately half the size of those in the Pseudoroegneria environments. Elymus growth could not be explained by simple resource competition alone; other mechanisms must have been involved in the apparent differences in interference patterns of neighboring plants with Elymus.

Human disturbance influences reproductive success and growth rate in California sea lions (Zalophus californianus)

The environment is currently undergoing changes at both global (e.g., climate change) and local (e.g., tourism, pollution, habitat modification) scales that have the capacity to affect the viability of animal and plant populations. Many of these changes, such as human disturbance, have an anthropogenic origin and therefore may be mitigated by management action. To do so requires an understanding of the impact of human activities and changing environmental conditions on population dynamics. We investigated the influence of human activity on important life history parameters (reproductive rate, and body condition, and growth rate of neonate pups) for California sea lions (Zalophus californianus) in the Gulf of California, Mexico. Increased human presence was associated with lower reproductive rates, which translated into reduced long-term population growth rates and suggested that human activities are a disturbance that could lead to population declines. We also observed higher body growth rates in pups with increased exposure to humans. Increased growth rates in pups may reflect a density dependent response to declining reproductive rates (e.g., decreased competition for resources). Our results highlight the potentially complex changes in life history parameters that may result from human disturbance, and their implication for population dynamics. We recommend careful monitoring of human activities in the Gulf of California and emphasize the importance of management strategies that explicitly consider the potential impact of human activities such as ecotourism on vertebrate populations.

Trade-off Between Plant Growth and Defense? A Comparison of Sagebrush populations

Abstract. We used ecotypic variation in big sagebrush (Artemisia tridentata) to examine potential trade-offs between inherent growth rate and tolerance or resistance to herbivory. Seeds were obtained from seven geographic populations, and 1,120 seedlings were established in a common garden. In one set of plots, plants were subjected to five treatments: control, regular insecticide spray, moderate browsing, severe browsing, or moderate browsing plus insecticide. Plants in a second set of plots were all untreated, and were used to estimate ambient growth, flower production, and susceptibility to herbivorous insects. In the first growing season, population differences in relative growth rate produced approximately seven-fold variation in mean biomass. Two populations of basin big sagebrush (A. tridentata tridentata) and one population of mountain big sagebrush (A. tridentata vaseyana) grew fastest; those of Wyoming big sagebrush (A. tridentata wyomingensis) showed the slowest growth. Bi-weekly application of insecticide for two growing seasons had no effect on the growth of either browsed or unbrowsed plants. All populations showed compensatory growth (but not overcompensation) in response to browsing, but the degree of compensation was unrelated to inherent growth rate. Similarly, there was no consistent relationship between plant growth rate and flower production in the second growing season. Some insects colonized fast-growing populations more frequently than slow-growing ones, but patterns of insect colonization were species-specific. At the level of geographic populations and subspecies, we found little evidence of a built-in trade-off between inherent growth rate and the ability to tolerate or resist herbivory. Because population ranks for growth rate changed substantially between seasons, attempts to correlate growth and defense characters need to account for differences in the growth trajectories of perennial plants.

Food restriction and chronic stress alter energy use and affect immunity in an infrequent feeder

Summary Glucocorticoids are important mediators of energy utilization for key physiological processes, including immune function. Much work has focused on the effects of energy limitation and stress for key physiological processes such as reproduction and immunity. However, it is unclear how stress alters energy use across different energy states, and the physiological ramifications of such effects are even less clear. In this study, we altered energy and stress states of an infrequent feeder, the terrestrial gartersnake (Thamnophis elegans), using fasting and repeated restraint stress (chronic stressors) to test how these challenges interacted to affect immune function, energy metabolites and glucocorticoid reactivity (a traditional indicator of stress state) to restraint stress, a standardized, acute stressor. After this acute stressor, the snakes which had received chronic stress had increased glucocorticoid reactivity, and both treatments altered energy metabolite use and storage. Evidence of interaction of food restriction and chronic stress treatments on innate immune function and energy metabolites (triglycerides and glycerol) suggests that stress alters energy use in a manner dependent on the energy state of the animal. Snakes have a remarkable ability to maintain functionality of key physiological processes under stressful conditions but are still susceptible to multiple simultaneous stressors, a situation increasingly prevalent in our ever-changing environment.

Effects of Rodent Species, Seed Species, and Predator Cues on Seed Fate

Abstract Seed selection, removal and subsequent management by granivorous animals is thought to be a complex interaction of factors including qualities of the seeds themselves (e.g., seed size, nutritional quality) and features of the local habitat (e.g. perceived predator risk). At the same time, differential seed selection and dispersal is thought to have profound effects on seed fate and potentially vegetation dynamics. In a feeding arena, we tested whether rodent species, seed species, and indirect and direct predation cues influence seed selection and handling behaviors (e.g., scatter hoarding versus larder hoarding) of two heteromyid rodents, Ord’s kangaroo rat (Dipodomys ordii) and the Great Basin pocket mouse (Perognathus parvus). The indirect cue was shrub cover, a feature of the environment. Direct cues, presented individually, were (1) control, (2) coyote (Canis latrans) vocalization, (3) coyote scent, (4) red fox (Vulpes vulpes) scent, or (5) short-eared owl (Asio flammeus) vocalization. We offered seeds of three sizes: two native grasses, Indian ricegrass (Achnatherum hymenoides) and bluebunch wheatgrass (Pseudoroegneria spicata), and the non-native cereal rye (Secale cereale), each in separate trays. Kangaroo rats preferentially harvested Indian ricegrass while pocket mice predominately harvested Indian ricegrass and cereal rye. Pocket mice were more likely to scatter hoard preferred seeds, whereas kangaroo rats mostly consumed and/or larder hoarded preferred seeds. No predator cue significantly affected seed preferences. However, both species altered seed handling behavior in response to direct predation cues by leaving more seeds available in the seed pool, though they responded to different predator cues. If these results translate to natural dynamics on the landscape, the two rodents are expected to have different impacts on seed survival and plant recruitment via their different seed selection and seed handling behaviors.

Competition on the range: science vs, perception in a bison-cattle conflict in the western USA

Summary Competition between livestock and wild ungulates is commonly perceived to occur on shared rangelands. In the Henry Mountains (HM) of Utah, a free‐ranging population of bison Bison bison has raised concerns among ranchers holding grazing permits on these public lands. Bison are the most conspicuous potential competitors with cattle, but lagomorphs (mainly jackrabbits Lepus californicus) are also abundant in this area. The local ranching community is applying political pressure on state and federal agencies to resolve ‘the bison problem’, but the relative grazing impacts of bison, cattle and lagomorphs have not previously been quantified. We constructed 40 grazing exclosures (each 5·95 m2) in the conflict area: 20 excluded bison + cattle (‘partial’) and 20 excluded bison + cattle + lagomorphs (‘full’). All exclosures, each with a paired open reference plot, were monitored for 1 year, and above‐ground plant production was measured. GPS telemetry (bison) and scheduled grazing (cattle) allowed visitation to be quantified for each ungulate species based on the number of ‘animal days’ in the area. Rancher perceptions of wildlife–cattle interactions were recorded in a questionnaire survey. Ranchers perceived bison as a high‐level competitor with cattle, whereas lagomorphs were perceived as low‐level competitors. Grazed reference plots yielded an average (±SE) of 22·7 g m−2 (±5·16) of grass, compared to 36·5 g m−2 (±7·33) in the partial exclosures and 43·7 g m−2 (±7·61) in the full exclosures. Exclusion of large herbivores thus resulted in a 13·8 g m−2 increase in grass biomass relative to the reference plots (P = 0·005), with the additional exclusion of lagomorphs resulting in a further 7·18 g m−2 increase (P = 0·048). Overall, lagomorphs accounted for 34·1%, bison 13·7% and cattle 52·3% of the total grass biomass removed by all herbivores on the shared range. Synthesis and applications. Cattle face a greater competitive challenge from lagomorphs than from bison in the study area. This case study illustrates the need for science‐based management of social–ecological systems in which even long‐term resource users might underestimate the complexities of trophic interactions. Attention should be redirected at the lagomorphs and their main predators, coyotes Canis latrans, which are currently subject to population control. To reduce negative perceptions among local ranchers, options should be explored to incorporate benefit‐sharing into the management of the bison population.

Dispersal behaviour of a polygynous carnivore: do cougars Puma concolor follow source-sink predictions?

The source-sink model of population dynamics predicts that density drives emigration of subordinate animals to habitats offering lower competition for resources. Several authors have suggested use of this model as a potential framework for conservation of exploited carnivores when precise enumeration is unfeasible. Dispersal is a critical behavioural mechanism for management based on this model, yet there is a lack of knowledge on the habitat and social conditions that motivate carnivore emigration and settlement. The cougar Puma concolor is a widely distributed and heavily exploited carnivore, indigenous to the western hemisphere. We evaluated patterns in cougar dispersal behaviour from two sites in Utah, differing in terms of management and the level of natural and anthropogenic habitat fragmentation. We used our results to evaluate three predictions with respect to cougar dispersal behaviour: 1) natal population density and maternal reproductive status prompt emigration, 2) movement of dispersing cougars is shaped by habitat configuration and permeability, and 3) dispersers preferentially settle in areas of high habitat quality and low conspecific density. We documented the emigration of 62 individuals and measured movement variables, including sex and site-specific frequency, distance, seasonality, direction and the habitat quality and harvest rates characterizing areas where immigrants settled. Although males and females exhibited pronounced differences in dispersal frequency, we found few differences in distance traveled, season of departure and direction moved. Dispersal occurred most frequently during spring, coinciding with the estrus pulse. Natural and anthropogenic obstacles modified landscape permeability, and therefore dispersal distances were shorter in fragmented habitats than in contiguous ones. Relative to males, females dispersed into habitats of lower productivity with higher mean annual harvest rates. Patterns in male settlement suggested habitat selection based on mating opportunities, whereas female settlement was predicated on avoiding conspecifics. Cougars in this Great Basin ecosystem largely conformed to source-sink predictions. Results can be used to parameterize source-sink models based on animal behaviour and landscape permeability to conserve exploited carnivores, under conditions of population expansion or recolonization of habitats where Allee effects are a limiting factor.