Bet hedging in a guild of desert annuals

Evolutionary bet hedging encapsulates the counterintuitive idea that organisms evolve traits that reduce short-term reproductive success in favor of longer-term risk reduction. It has been widely investigated theoretically, and many putative examples have been cited including practical ones such as the dormancy involved in microbe and weed persistence. However, long-term data on demographic variation from the actual evolutionarily relevant environments have been unavailable to test for its mechanistic relationship to alleged bet hedging traits. I report an association between delayed germination (a bet hedging trait) and risk using a 22-year data set on demographic variation for 10 species of desert annual plants. Species with greater variation in reproductive success (per capita survival from germination to reproduction x per capita fecundity of survivors) were found to have lower average germination fractions. This provides a definitive test using realistic data on demographic variance that confirms the life history prediction for bet hedging. I also showed that the species with greater long-term demographic variation tended to be the ones with greater sensitivity of reproductive success to variation among years in growing-season precipitation.     

New methods for quantifying the spatial storage effect: an illustration with desert annuals

Recent theory supports the long-held proposition that coexistence is promoted by species-specific responses to a spatially varying environment. The underlying coexistence mechanism, the spatial storage effect, can be quantified by the covariance between response to the environment and competition. Here, "competition" is generalized to encompass similar processes such as facilitation and apparent competition. In the present study, we use a model field system of desert annual plants to demonstrate this method and to provide insight into the dynamics of the field system. Specifically, we use neighborhood competition experiments to quantify the spatial storage effect and compare it to the separate (but not mutually exclusive) process of neighborhood-scale resource partitioning. As our basic experimental design has been used frequently in community ecology, these methods can be applied to many existing data sets, as well as future field studies.     

Seed dispersal by pulp consumers, not "legitimate" seed dispersers, increases Guettarda viburnoides population growth

We examined the effect of seed dispersal by Purplish Jays (Cyanocorax cyanomelas; pulp consumers) and the Chestnut-eared Ara?ari (Pteroglossus castanotis; "legitimate" seed dispersers) on population growth of the small tree Guettarda viburnoides (Rubiaceae) in northeastern Bolivian savannas. Because each bird species differs with respect to feeding and post-feeding behavior, we hypothesized that seed dispersal by each species will contribute differently to the rate of increase of G. viburnoides, but that seed dispersal by either species will increase population growth when compared to a scenario with no seed dispersal. To examine the effects of individual dispersers on the future population size of G. viburnoides, we projected population growth rate using demographic models for G. viburnoides that explicitly incorporate data on quantitative and qualitative aspects of seed dispersal by each frugivore species. Our model suggests that seed dispersal by C. cyanomelas leads to positive population growth of G. viburnoides, whereas seed dispersal by P. castanotis has a detrimental effect on the population growth of this species. To our knowledge, this is the first study to report negative effects of a "legitimate" seed disperser on the population dynamics of the plant it consumes. Our results stress the importance of incorporating frugivore effects into population projection matrices, to allow a comprehensive analysis of the effectiveness of different dispersers for plant population dynamics.     

Explaining long-distance dispersal: effects of dispersal distance on survival and growth in a stream salamander

Long-distance dispersal (LDD) may contribute disproportionately to range expansions, the creation of new evolutionary lineages, and species persistence in human-dominated landscapes. However, because data on the individual consequences of dispersal distance are extremely limited, we have little insight on how LDD is maintained in natural populations. I used six years of spatially explicit capture-mark-recapture (CMR) data to test the prediction that individual performance increases with dispersal distance in the stream salamander Gyrinophilus porphyriticus. Dispersal distance was total distance moved along the 1-km study stream, ranging from 0 to 565 m. To quantify individual performance, I used CMR estimates of survival and individual growth rates based on change in body length. Survival and growth rates increased significantly with dispersal distance. These relationships were not confounded by pre-dispersal body condition or by ecological gradients along the stream. Individual benefits of LDD were likely caused by an increase in the upper limit of settlement site quality with dispersal distance. My results do not support the view that the fitness consequences of LDD are unpredictable and instead suggest that consistent evolutionary mechanisms may explain the prevalence of LDD in nature. They also highlight the value of direct CMR data for understanding the individual consequences of variation in dispersal distance and how that variation is maintained in natural populations.     

Assessing biodegradation benefits from dispersal networks

The performance of biodegradation of organic pollutants in soil often depends on abiotic conditions and the bioavailability of these pollutants to degrading bacteria. In this context, bacterial dispersal is an essential aspect. Recent studies on the potential promotion of bacterial dispersal by fungal hyphae raised the idea of specifically applying fungal networks to accelerate bacterial degradation processes in situ. Our objective is to investigate these processes and their performance via simulation modelling and address the following questions: (1) Under what abiotic conditions can dispersal networks significantly improve bacterial degradation? and (2) To what extent does the spatial configuration of the networks influence the degradation performance? To answer these questions, we developed a spatially explicit bacterial colony model, which is applied to controlled laboratory experiments with Pseudomonas putida G7 organisms as a case study. Using this model, we analyzed degradation performance in response to different environmental scenarios and showed that conditions of limited bacterial dispersal also limit degradation performance. Under such conditions, dispersal networks have the highest potential for improving the bioavailability of pollutants to bacteria. We also found that degradation performance significantly varies with the spatial configuration of the dispersal networks applied and the time horizon over which performance is assessed. Regarding future practical applications, our results suggest that (1) fungal networks may dramatically improve initially adverse conditions for biodegradation of pollutants in soil, and (2) the network's spatial structure and accessibility are decisive for the success of such tasks.     

Episodic death across species of desert shrubs

Extreme events shape population and community trajectories. We report episodic mortality across common species of thousands of long-lived perennials individually tagged and monitored for 20 years in the Colorado Desert of California following severe regional drought. Demographic records from 1984 to 2004 show 15 years of virtual stasis in populations of adult shrubs and cacti, punctuated by a 55-100% die-off of six of the seven most common perennial species. In this episode, adults that experienced reduced growth in a lesser drought during 1984-1989 failed to survive the drought of 2002. The significance of this event is potentially profound because population dynamics of long-lived plants can be far more strongly affected by deaths of adults, which in deserts potentially live for centuries, than by seedling births or deaths. Differential mortality and rates of recovery during and after extreme climatic events quite likely determine the species composition of plant and associated animal communities for at least decades. The die-off recorded in this closely monitored community provides a unique window into the mechanics of this process of species decline and replacement.     

Bright moonlight triggers natal dispersal departures

Upon leaving their natal area, dispersers are confronted with unknown terrains. Species-specific perceptual ranges (i.e. the maximum distance from which an individual can perceive landscape features) play a crucial role in spatial movement decisions during such wanderings. In nocturnal animals that rely on vision, perceptual range is dramatically enhanced during moonlight, compared to moonless conditions. This increase of the perceptual range is an overlooked element that may be responsible for the successful crossing of unfamiliar areas during dispersal. The information gathered from 143 radio-tagged eagle owl Bubo bubo juveniles in Spain, Finland and Switzerland shows that, although the decision to initiate dispersal is mainly an endogenous phenomenon determined by the attainment of a given age (～6 months), dispersers leave their birthplace primarily under the best light conditions at night, i.e. when most of the lunar disc is illuminated. This sheds new light into the mechanisms that may trigger dispersal from parental territory.     

Analyzing maps of dispersal around a single focus

The spatial pattern of organisms may be used to characterize their dispersal, quantify spread or estimate the point of introduction of an alien species. Their distribution may be represented by maps of individuals, or by counts or by presence/absence at known positions within a sampled area. The problems and relative merits of these different forms of data for spatial inference are discussed. Three datasets concerning dispersal from a single focus are analyzed: counts of aphids, Rhopalosiphum padi and Sitobion avenae, on barley plants, Hordeum vulgare, grown in experi- mental trays; mapped locations of couch grass, Elymus repens, tillers within plots of a field experiment; locations of sightings of the lupin aphid, Macrosiphum albifrons, as it invaded Great Britain between 1981 and 1984. A method for generating maps from counts is proposed to overcome problems caused by recording imprecision. Several statistics are used to quantify dispersal and spatial pattern in the experimental data and together provide a clear picture of the spatial pattern observed; they enabled several effects of the experimental treatments to be identified. The value of the statistics are compared. Estimates of the source of the lupin aphid invasion are obtained using the backtracking methods of Perry (1995b) and do not contradict previous suggestions.     

Sex-specific fitness consequences of dispersal in Siberian jays

In most birds, natal dispersal is female-biased, but the selective pressures leading to this pattern have rarely been explored with comprehensive data on lifetime reproductive success. In territorial birds, the benefit of philopatry should be higher for males than for females when males establish territories for which knowledge about the local environment is important. As females may use male characteristics for mate choice, and hence indirectly for territory choice, the benefit from the direct knowledge of the local environment may be lower for females than males. We tested this hypothesis using data from a long-term study of group living corvids, the Siberian jays (Perisoreus infaustus). In this species, the socially dominant offspring delay dispersal while the sub-dominant offspring leave the family group directly after reaching independence. Our results show that natal dispersal distance (a proxy for local knowledge) was related to sex and dispersal timing (a proxy for “quality”): Females and early dispersers traveled further on average than males and delayed dispersers. Furthermore, dispersal distance and timing were negatively related to the number of recruits produced over an individual’s lifetime in males, but not in females. Hence, the results support the hypothesis that the female-biased natal dispersal found in this and other bird species may come about through higher lifetime reproductive success of philopatric males than females.     

Influence of maternal food availability on offspring dispersal

Prenatal effects caused by the maternal environment during gestation are known to contribute to the phenotype of the offspring. Whether they have some adaptive value is currently under debate. We experimentally tested the existence of such a maternal effect (food availability during gestation) on dispersal of offspring in the common lizard (Lacerta vivipara). Pregnant females were captured and kept in the laboratory until parturition. During this period, females were offered two rates of food delivery. After parturition, we released mothers and offspring at the mother's capture point. Dispersal of young was significantly affected by the mother's nutrition. To our knowledge, this is the first evidence of a prenatal effect on dispersal. Offspring of well fed mothers dispersed at a higher rate than those of less well fed mothers. As current hypotheses clearly predict the opposite result, our evidence calls for their reassessment. Dispersers are not always the least fit individuals or those coming from the poorest environments.     

荒漠河岸生态系统退化机理的定量分析

通过野外监测资料,采用聚类分析与主成分分析法,分析了荒漠河岸生态系统退化的内外因素与驱动力、植被退化的环境主导因子与生态退化程度,结果表明:(1)地质、地貌与气候条件等因素的影响,塔里木河下游生态系统存在脆弱和不稳定性是其退化的内因,外界干扰则是其退化的外在因素。人为干扰则是其退化发生的驱动力,起因于人口的增加、需求的增长。(2)在此过程中导致植被受损的环境主导因子是水,植被退化是人类干扰作用于植被赖以生存的环境主导因子所致。(3)其不同河段的生态退化可以归为三类:潜在沙漠化类(轻度退化),轻度沙漠化类(中度退化),中度或重度沙漠化类(重度退化),其中第三类型,土壤条件与其它两类差异较大,表现出土壤退化的滞后现象。     

Increasing accuracy of dispersal kernels in grid-based population models

Dispersal kernels in grid-based population models specify the proportion, distance and direction of movements within the model landscape. Spatial errors in dispersal kernels can have large compounding effects on model accuracy. Circular Gaussian and Laplacian dispersal kernels at a range of spatial resolutions were investigated, and methods for minimizing errors caused by the discretizing process were explored. Kernels of progressively smaller sizes relative to the landscape grid size were calculated using cell-integration and cell-center methods. These kernels were convolved repeatedly, and the final distribution was compared with a reference analytical solution. For large Gaussian kernels (σ > 10 cells), the total kernel error was <10⁻¹¹ compared to analytical results. Using an invasion model that tracked the time a population took to reach a defined goal, the discrete model results were comparable to the analytical reference. With Gaussian kernels that had σ ≤ 0.12 using the cell integration method, or σ ≤ 0.22 using the cell center method, the kernel error was greater than 10%, which resulted in invasion times that were orders of magnitude different than theoretical results. A goal-seeking routine was developed to adjust the kernels to minimize overall error. With this, corrections for small kernels were found that decreased overall kernel error to <10⁻¹¹ and invasion time error to <5%.     

荒漠河岸生态系统退化机理的定量分析

通过野外监测资料,采用聚类分析与主成分分析法,分析了荒漠河岸生态系统退化的内外因素与驱动力、植被退化的环境主导因子与生态退化程度,结果表明：（1）地质、地貌与气候条件等因素的影响,塔里木河下游生态系统存在脆弱和不稳定性是其退化的内因,外界干扰则是其退化的外在因素。人为干扰则是其退化发生的驱动力,起因于人口的增加、需求的增长。（2）在此过程中导致植被受损的环境主导因子是水,植被退化是人类干扰作用于植被赖以生存的环境主导因子所致。（3）其不同河段的生态退化可以归为三类：潜在沙漠化类（轻度退化）,轻度沙漠化类（中度退化）,中度或重度沙漠化类（重度退化）,其中第三类型,土壤条件与其它两类差异较大,表现出土壤退化的滞后现象。     

Habitat loss through disruption of constrained dispersal networks.

Large losses of habitat could be caused by land use change that disrupts the dispersal networks used by migratory species. We assessed the relative losses of habitat for diadromous fish (i.e., those migrating between sea and freshwater) due to physical barriers, degradation of migratory passage associated with catchment land use, and site-scale land use characteristics on the West Coast, South Island, New Zealand. Fish occurrence, land use data, and river network models were analyzed in a GIS and subjected to a three-level hierarchical analysis. To identify accessible habitat not restricted by physical barriers, we used the migratory distance and maximum downstream slope encountered to identify accessible sites in least-impacted catchments and applied the results to all catchments within the study area. For two fish species, banded kokopu (Galaxias fasciatus) and koaro (G. brevipinnis), sites modeled as accessible using logistic regression in least-impacted catchments were then used to assess the impacts of catchment-scale deforestation and downstream land uses on habitat loss. Finally, sites not restricted by physical barriers or land-use-related impacts on migratory passage were used to model the effects of local land use. The models indicated that koaro and banded kokopu potentially had access to 28,000 km and 5300 km, respectively, of the 40,600 km of streams within the study area. Impacts due to intensive agricultural land use downstream in catchments affecting migratory passage were predicted to reduce the accessible habitats for koaro and banded kokopu by 55% and 70%, respectively. Local land use further reduced koaro and banded kokopu habitats to 70% and 90%, respectively, of total accessible habitat. Habitat lost through disruption of the dispersal network was disproportionately large because potentially useable habitat was rendered inaccessible.     

Can non-breeding be a cost of breeding dispersal?

Breeding habitat selection and dispersal are crucial processes that affect many components of fitness. Breeding dispersal entails costs, one of which has been neglected: dispersing animals may miss breeding opportunities because breeding dispersal requires finding a new nesting site and mate, two time- and energy-consuming activities. Dispersers are expected to be prone to non-breeding. We used the kittiwake (Rissa tridactyla) to test whether breeding dispersal influences breeding probability. Breeding probability was associated with dispersal, in that both were negatively influenced by private information (previous individual reproductive success) and public information (average reproductive success of conspecifics) about patch quality. Furthermore, the probability of skipping breeding was 1.7 times higher in birds that settled in a new patch relative to those that remained on the same patch. Finally, non-breeders that resumed breeding were 4.4 times more likely to disperse than birds that bred in successive years. Although private information may influence breeding probability directly, the link between breeding probability and public information may be indirect, through the influence of public information on breeding dispersal, non-breeding thus being a cost of dispersal. These results support the hypothesis that dispersal may result in not being able to breed. More generally, non-breeding (which can be interpreted as an extreme form of breeding failure) may reveal costs of various previous activities. Because monitoring the non-breeding portion of a population is difficult, non-breeders have been neglected in many studies of reproduction trade-offs.     

Ontogeny of dispersal distances in young Spanish imperial eagles

Summary I studied factors influencing dispersal distances in 30 young Spanish imperial eagles (Aquila adalberti) radio-tagged in southwestern Spain in 1986–1990. The mean dispersal distance between the natal nest and the settling area was 138 km. Every young bird reached its maximum dispersal distance within 4 months of its departure from the natal population. No significant differences between the sexes were detected in maximum dispersal distance, but females spent more time in the more distant settling areas. Dispersal distance was not related to the date of departure from the natal population. Dispersal distance was significantly longer for birds that hatched earlier and that had lower blood urea levels, indicating that better-nourished young had longer displacements. These results are not in accordance with the competitive displacement hypothesis.     

Changes of movement patterns from early dispersal to settlement

Moving and spatial learning are two intertwined processes: (a) changes in movement behavior determine the learning of the spatial environment, and (b) information plays a crucial role in several animal decision-making processes like movement decisions. A useful way to explore the interactions between movement decisions and learning of the spatial environment is by comparing individual behaviors during the different phases of natal dispersal (when individuals move across more or less unknown habitats) with movements and choices of breeders (who repeatedly move within fixed home ranges), that is, by comparing behaviors between individuals who are still acquiring information vs. individuals with a more complete knowledge of their surroundings. When analyzing movement patterns of eagle owls, Bubo bubo, belonging to three status classes (floaters wandering across unknown environments, floaters already settled in temporary settlement areas, and territory owners with a well-established home range), we found that: (1) wandering individuals move faster than when established in a more stable or fixed settlement area, traveling larger and straighter paths with longer move steps; and (2) when floaters settle in a permanent area, then they show movement behavior similar to territory owners. Thus, movement patterns show a transition from exploratory strategies, when animals have incomplete environmental information, to a more familiar way to exploit their activity areas as they get to know the environment better.     

Precipitation-driven carbon balance controls survivorship of desert biocrust mosses

Precipitation patterns including the magnitude, timing, and seasonality of rainfall are predicted to undergo substantial alterations in arid regions in the future, and desert organisms may be more responsive to such changes than to shifts in only mean annual rainfall. Soil biocrust communities (consisting of cyanobacteria, lichen, and mosses) are ubiquitous to desert ecosystems, play an array of ecological roles, and display a strong sensitivity to environmental changes. Crust mosses are particularly responsive to changes in precipitation and exhibit rapid declines in biomass and mortality following the addition of small rainfall events. Further, loss of the moss component in biocrusts leads to declines in crust structure and function. In this study, we sought to understand the physiological responses of the widespread and often dominant biocrust moss Syntrichia caninervis to alterations in rainfall. Moss samples were collected during all four seasons and exposed to two rainfall event sizes and three desiccation period (DP) lengths. A carbon balance approach based on single precipitation events was used to define the carbon gain or loss during a particular hydration period. Rainfall event size was the strongest predictor of carbon balance, and the largest carbon gains were associated with the largest precipitation events. In contrast, small precipitation events resulted in carbon deficits for S. caninervis. Increasing the length of the DP prior to an event resulted in reductions in carbon balance, probably because of the increased energetic cost of hydration following more intense bouts of desiccation. The season of collection (i.e., physiological status of the moss) modulated these responses, and the effects of DP and rainfall on carbon balance were different in magnitude (and often in sign) for different seasons. In particular, S. caninervis displayed higher carbon balances in the winter than in the summer, even for events of identical size. Overall, our results suggest that annual carbon balance and survivorship in biocrust mosses are largely driven by precipitation, and because of the role mosses play in biocrusts, changes in intra-annual precipitation patterns can have implications for hydrology, soil stability, and nutrient cycling in dryland systems.