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.     

Density dependence vs. independence, and irregular population dynamics of a swallow-wort fruit fly

Although most long-term studies of consumer-resource (e.g., predator-prey) interactions select species showing cyclic population dynamics, strong consumer-resource interactions can also produce irregular, noncyclic dynamics. Here, we present a case in which a seed predator, the tephritid fruit fly Euphranta connexa, shows fluctuations in density of more than two orders of magnitude over a 22-year period. To explain these fluctuations, we analyzed a stage-specific data set to quantify the density-dependent and density-independent components of larval survivorship and realized fecundity. Both larval survivorship and realized fecundity were strongly density dependent. Larval survivorship dropped from 0.62 at low larval density to 0.081 at high larval density, whereas fecundity dropped from 84.3 to 0.32 eggs per individual, more than a 100-fold decrease. We divided density-independent variation in E. connexa population dynamics into components for variability in (1) larval survivorship, (2) realized fecundity, and (3) annual fruit abundance. Of these components, 96% of the density-independent variance in per capita population growth rates was caused by fluctuations in fruit abundance. This highlights the importance of the strong consumer-resource interactions in driving fluctuations in E. connexa abundance. It also demonstrates that E. connexa dynamics are remarkably simple, and aside from the 4% of unexplained variance in per capita population growth rates, our understanding of E. connexa dynamics is remarkably complete.     

Relatedness and environment affect traits associated with invasive and noninvasive introduced Commelinaceae.

Understanding the traits of invasive species may improve the ability to predict, prevent, and manage invasions. I compared morphological and performance traits of five congeneric pairs of invasive and noninvasive Commelinaceae across a factorial experiment using a range of water and nutrient availabilities. Invasive species had greater fecundity and vegetative reproduction than their noninvasive relatives. The invasive species also had higher relative growth rates, greater specific leaf area, and more plastic root-to-shoot ratios than noninvasive species. However, whether a trait was associated with invasiveness often depended on both environment and relatedness. Invasives had greater sexual and vegetative reproduction, higher specific leaf area, and greater relative growth rates than noninvasive congeners, but only in some environments. Differences between invasive and noninvasive taxa were greatest at high nutrient availabilities. These results suggest that studies of invasive species' traits must incorporate information on conditions under which the trait was measured. In addition, incorporating information on relatedness improved our ability to detect associations between species traits, such as specific leaf area and relative growth rate, and invasiveness, suggesting that such information may be required for a complete understanding of what makes a species invasive.     

Stochastic population dynamics in populations of western terrestrial garter snakes with divergent life histories

Comparative evaluations of population dynamics in species with temporal and spatial variation in life-history traits are rare because they require long-term demographic time series from multiple populations. We present such an analysis using demographic data collected during the interval 1978-1996 for six populations of western terrestrial garter snakes (Thamnophis elegans) from two evolutionarily divergent ecotypes. Three replicate populations from a slow-living ecotype, found in mountain meadows of northeastern California, were characterized by individuals that develop slowly, mature late, reproduce infrequently with small reproductive effort, and live longer than individuals of three populations of a fast-living ecotype found at lakeshore locales. We constructed matrix population models for each of the populations based on 8-13 years of data per population and analyzed both deterministic dynamics based on mean annual vital rates and stochastic dynamics incorporating annual variation in vital rates. (1) Contributions of highly variable vital rates to fitness (lambda(s)) were buffered against the negative effects of stochastic variation, and this relationship was consistent with differences between the meadow (M-slow) and lakeshore (L-fast) ecotypes. (2) Annual variation in the proportion of gravid females had the greatest negative effect among all vital rates on lambda(s). The magnitude of variation in the proportion of gravid females and its effect on lambda(s) was greater in M-slow than L-fast populations. (3) Variation in the proportion of gravid females, in turn, depended on annual variation in prey availability, and its effect on lambda(s) was 4 23 times greater in M-slow than L-fast populations. In addition to differences in stochastic dynamics between ecotypes, we also found higher mean mortality rates across all age classes in the L-fast populations. Our results suggest that both deterministic and stochastic selective forces have affected the evolution of divergent life-history traits in the two ecotypes, which, in turn, affect population dynamics. M-slow populations have evolved life-history traits that buffer fitness against direct effects of variation in reproduction and that spread lifetime reproduction across a greater number of reproductive bouts. These results highlight the importance of long-term demographic and environmental monitoring and of incorporating temporal dynamics into empirical studies of life-history evolution.     

Extending the leaf economics spectrum to decomposition: evidence from a tropical forest

I investigated the relationship between leaf physiological traits and decomposition of leaf litter for 35 plant species of contrasting growth forms from a lowland tropical forest in Panama to determine whether leaf traits could be used to predict decomposition. Decomposition rate (k) was correlated with specific leaf area (SLA), leaf nitrogen (N), phosphorus (P), and potassium (K) across all species. Photosynthetic rate per unit mass (Amass) was not correlated with k, but structural equation modeling showed support for a causal model with significant indirect effects of Amass on k through SLA, N, and P, but not K. The results indicate that the decomposability of leaf tissue in this tropical forest is related to a global spectrum of leaf economics that varies from thin, easily decomposable leaves with high nutrient concentrations and high photosynthetic rates to thick, relatively recalcitrant leaves with greater physical toughness and defenses and low photosynthetic rates. If this pattern is robust across biomes, then selection for suites of traits that maximize photosynthetic carbon gain over the lifetime of the leaf may be used to predict the effects of plant species on leaf litter decomposition, thus placing the ecosystem process of decomposition in an evolutionary context.     

Size-mediated non-trophic interactions and stochastic predation drive assembly and dynamics in a seabird community

Theoretical and empirical evidence suggests that body size is a major life-history trait impacting on the structure and functioning of complex food webs. However, long-term analyses of size-dependent interactions within simpler network modules, for instance, competitive guilds, are scant. Here, we model the assembly dynamics of the largest breeding seabird community in the Mediterranean basin during the last 30 years. This unique data set allowed us to test, through a "natural experiment," whether body size drove the assembly and dynamics of an ecological guild growing from very low numbers after habitat protection. Although environmental stochasticity accounted for most of community variability, the population variance explained by interspecific interactions, albeit small, decreased sharply with increasing body size. Since we found a demographic gradient along a body size continuum, in which population density and stability increase with increasing body size, the numerical effects of interspecific interactions were proportionally higher on smaller species than on larger ones. Moreover, we found that the per capita interaction coefficients were larger the higher the size ratio among competing species, but only for the set of interactions in which the species exerting the effect was greater. This provides empirical evidence for long-term asymmetric interspecific competition, which ultimately prompted the local extinction of two small species during the study period. During the assembly process stochastic predation by generalist carnivores further triggered community reorganizations and global decays in population synchrony, which disrupted the pattern of interspecific interactions. These results suggest that the major patterns detected in complex food webs can hold as well for simpler sub-modules of these networks involving non-trophic interactions, and highlight the shifting ecological processes impacting on assembling vs. asymptotic communities.     

Bridging the generation gap in plants: pollination, parental fecundity, and offspring demography

Despite extensive study of pollination and plant reproduction on the one hand, and of plant demography on the other, we know remarkably little about links between seed production in successive generations, and hence about long-term population consequences of variation in pollination success. We bridged this "generation gap" in Ipomopsis aggregata, a long-lived semelparous wildflower that is pollinator limited, by adding varying densities of seeds to natural populations and following resulting plants through their entire life histories. To determine whether pollen limitation of seed production constrains rate of population growth in this species, we sowed seeds into replicated plots at a density that mimics typical pollination success and spacing of flowering plants in nature, and at twice that density to mimic full pollination. Per capita offspring survival, flower production, and contribution to population increase (lambda) did not decline with sowing density in this experiment, suggesting that typical I. aggregata populations freed from pollen limitation will grow over the short term. In a second experiment we addressed whether density dependence would eventually erase the growth benefits of full pollination, by sowing a 10-fold range of seed densities that falls within extremes estimated for the natural "seed rain" that reaches the soil surface. Per capita survival to flowering and age at flowering were again unaffected by sowing density, but offspring size, per capita flower production, and lambda declined with density. Such density dependence complicates efforts to predict population dynamics over the longer term, because it changes components of the life history (in this case fecundity) as a population grows. A complete understanding of how constraints on seed production affect long-term population growth will hinge on following offspring fates at least through flowering of the first offspring generation, and doing so for a realistic range of population densities.     

宁夏六盘山8种木本植被的叶片水分利用效率

为了解宁夏六盘山地区主要树种的叶片水分利用效率（WUE）的种间和水分生境差异及季节变化,2010年6-9月份在宁夏六盘山具有半湿润气候和较多树种的香水河小流域和具有半干旱气候及较少树种的叠叠沟小流域,选择8个主要树种测定其叶片的 WUE。结果表明,（1）在香水河小流域,树种叶片 WUE 大小排序为油松 Pinus tablaeformis＞山桃 Prunus davidiana＞沙棘Hippophae rhamnoides＞辽东栎Quercus liaotungensis＞华北落叶松Larix principris-rupprechtii＞华山松Pinus armandii＞少脉椴Tilia paucicostata＞白桦Betula platyphylla;在不同生活型之间,叶片WUE的大小排序基本上呈现出灌木＞小乔木＞常绿针叶树＞落叶针叶树＞落叶阔叶树的变化规律;所有8个测试树种的叶片WUE的季节变化均为生长季初期较高,在中后期持续降低,但季节变幅随树种而异。（2）对2个小流域共有的测试树种（华北落叶松、山桃和沙棘）的叶片WUE进行了比较,结果表明存在着明显的水分生境差异,即在半干旱区极显著（p＜0.01）大于半湿润区,前者为后者的2.03倍以上。（3）对于沙棘、山桃、油松和华北落叶松,它们在较干旱气候类型立地和较干旱季节条件下都具有较高的叶片WUE,属抗旱能力较强的生态型节水树种,可选为干旱缺水区主要造林树种。此外,树种叶片WUE同时具有保守性和变异性,说明可采取适度抗旱锻炼来在一定范围内提高造林树种的WUE。     

东江中上游主要造林树种光合生理特征

在东江中上游,以6年生造林树种木荷Schima superba、红锥Castanopsis hystrix、火力楠Michelia macclurei和藜蒴Castanopsis fissa为材料,利用Li-6400光合作用测定系统,对东江中上游水源林主要造林树种木荷、红锥、火力楠和藜蒴叶片光合日变化和光响应进行了测定,探讨了4种树种在当地自然生境下的光合生理特征。结果表明：木荷和藜蒴净光合速率（Pn）日变化呈＂单峰＂曲线,火力楠和红锥净光合速率（Pn）日变化呈＂双峰＂曲线,4种树种净光合速率（Pn）最大值均出现在10：00左右,分别为8.73、6.00、6.68和7.91μmol.m-1.s-1;4种树种净光合速率（Pn）日均值表现为木荷〉火力楠〉红锥〉藜蒴,蒸腾速率（Tr）日均值表现为红锥〉木荷〉火力楠〉藜蒴,水分利用效率（WUE）日均值表现为木荷〉藜蒴〉火力楠〉红锥,木荷具有高光合、高水分利用效率的生理特性,对环境适应能力强。4种树种光响应曲线均符合非直角双曲线模型（R2〉0.98）和指数模型（R2〉0.98）,采用非直角双曲线模型拟合的4种树种最大净光合速率（P,max）、暗呼吸速率（Rd）和表观量子效率（a）基本高于指数模型相应拟合值,且接近实测值,但2种模型拟合的光饱和点低于实际观测值。在中幼龄阶段,木荷、红锥、火力楠和藜蒴表现出较高的光饱和点（LSP）、较低的光补偿点（LCP）和较低表观量子效率,具有较强的耐阴喜光特性,对光照强度表现出一定的适应性和可塑性,对弱光与强光的利用能力较高。这些研究结果为东江流域水源林造林设计、树种筛选和抚育提供理论参考。     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

Individual and combined effects of Ca/Mg ratio and water on trait expression in Mimulus guttatus

Low Ca/Mg ratios (a defining component of serpentine soils) and low water environmental conditions often co-occur in nature and are thought to exert strong selection pressures on natural populations. However, few studies test the individual and combined effects of these environmental factors. We investigated the effects of low Ca/Mg ratio and low water availability on plant leaf, stem, stolon, and floral traits of Mimulus guttatus, a bodenvag species, i.e., a species that occurs in serpentine and non-serpentine areas. We quantified genetic variation and genetic variation for plasticity for these leaf, stem, stolon, and floral traits at three hierarchical levels: field-habitat type, population, and family, and we evaluated the relative importance of local adaptation and plasticity. We chose two populations and 10 families per population from four distinct field "habitat types" in northern California: high Ca/Mg ratio (non-serpentine) and season-long water availability, high Ca/Mg ratio and seasonally drying, low Ca/Mg ratio (serpentine) and season-long water availability, and low Ca/Mg ratio and seasonally drying. Seedlings were planted into greenhouse treatments that mimicked the four field conditions. We only detected genetic variation for stem diameter and length of longest leaf at the field-habitat level, but we detected genetic variation at the family level for nearly all traits. Soil chemistry and water availability had strong phenotypic effects, alone and in combination. Our hypothesis of an association between responses to low water levels and low Ca/Mg ratio was upheld for length of longest leaf, stem diameter, corolla width, and total number of reproductive units, whereas for other traits, responses to Ca/Mg ratio and low water were clearly independent. Our results suggest that traits may evolve independently from Ca/Mg ratios and water availability and that our focal traits were not simple alternative measures of vigor. We found genetic variation for plasticity both at the field-habitat type and family levels for half of the traits studied. Phenotypic plasticity and genetic variation for plasticity appear to be more important than local adaptation in the success of these M. guttatus populations found across a heterogeneous landscape in northern California. Phenotypic plasticity is an important mechanism maintaining the broad ecological breadth of native populations of M. guttatus.     

Ecology of hemiepiphytism in fig species is based on evolutionary correlation of hydraulics and carbon economy

Woody hemiepiphytic species (Hs) are important components of tropical rain forests, and they have been hypothesized to differ from non-hemiepiphytic tree species (NHs) in adaptations relating to water relations and carbon economy; but few studies have been conducted comparing ecophysiological traits between the two growth forms especially in an evolutionary context. Using common-garden plants of the genus Ficus, functional traits related to plant hydraulics and carbon economy were compared for seven NHs and seven Hs in their adult terrestrial "tree-like" growth phase. We used phylogenetically independent contrasts to test the hypothesis that differences in water availability selected for contrasting suites of traits in Hs and NHs, driving evolutionary correlations among functional traits including hydraulic conductivity and photosynthetic traits. Species of the two growth forms differed in functional traits; Hs had substantially lower xylem hydraulic conductivity and stomatal conductance, and higher instantaneous photosynthetic water use efficiency. Leaf morphological and structural traits also differed strikingly between the two growth forms. The Hs had significantly smaller leaves, higher leaf mass per area (LMA), and smaller xylem vessel lumen diameters. Across all the species, hydraulic conductivity was positively correlated with leaf gas exchange indicating high degrees of hydraulic-photosynthetic coordination. More importantly, these correlations were supported by correlations implemented on phylogenetic independent contrasts, suggesting that most trait correlations arose through repeated convergent evolution rather than as a result of chance events in the deep nodes of the lineage. Vatiation in xylem hydraulic conductivity was also centrally associated with a suite of other functional traits related to carbon economy and growth, such as LMA, water use efficiency, leaf nutrient concentration, and photosynthetic nutrient use efficiency, indicating important physiological constraints or trade-offs among functional traits. Shifts in this trait cluster apparently related to the adaptation to drought-prone canopy growth during the early life cycle of Hs and clearly affected ecophysiology of the later terrestrial stage of these species. Evolutionary flexibility in hydraulics and associated traits might be one basis for the hyper-diversification of Ficus species in tropical rain forests.     

A resource-depletion model of forest insect outbreaks

Detailed analyses of thresholded ecological interactions can improve our understanding of the transition from aperiodic to periodic dynamics. We develop a threshold model of the population dynamics of outbreaking bark beetle populations that alternate between non-epidemic and epidemic behavior. The model involves accumulation of resources during low-density periods and depletion during outbreaks. The transition between the two regimes is caused by disturbance events in the form of major tree felling by wind. The model is analyzed with particular reference to the population dynamics of the spruce bark beetle (Ips typographus) in Scandinavia for which a comprehensive literature allows full parameterization. The fairly constant outbreak lengths and the highly variable waiting time between outbreaks that are seen in the historical records of this species agree well with the predictions of the model. The thresholded resource-depletion dynamics result in substantial variation in the degree of periodicity between stochastic realizations. The completely aperiodic tree colonizations are partly predictable when the timing of the irregular windfall events are known. However, the predictability of inter-outbreak periods is low due to the large variation of cases falling most frequently in the middle between the extremes of purely nonperiodic (erratic) and periodic (cyclic) fluctuations.     

Population Size and Reproduction in Phlox pilosa

Abstract: We analyzed the relationships between population size and reproductive characteristics in the perennial prairie forb Phlox pilosa , an obligate outcrossing butterfly-pollinated species. We examined 27 populations ranging in size from 9 to over 75,000 flowering ramets in two regions of the state of Iowa (eastcentral and northwest) in 1993 and 1994. We collected flowers from each population and scored them for pollen arrival to stigmas and number of pollen tubes. We collected fruiting ramets from each population at the end of the Phlox growing season and scored them for height, biomass, and reproductive variables, including the number of flowers initiated and opened and the number of capsules initiated and matured. In both years, population size was significantly correlated with the number of capsules matured per ramet. Differences between populations in capsule production were set primarily at the pollination stage. In 1993, pollen arrival to stigmas was significantly lower than in 1994 and was correlated with population size in eastcentral Iowa populations, indicating that lower reproduction in small populations that year was at least partially due to inadequate amounts of pollen being moved. In 1993, weather conditions likely depressed pollinator activity, but absolute capsule formation was high because of high flower production per ramet and high population densities. In 1994, when pollen arrival to stigmas was relatively high and unrelated to population size, outcross pollen movement was greater in larger populations. Increased efficacy of outcross pollen movement in 1994 may have resulted from lower flower production and less dense populations forcing greater pollinator movement between ramets or from variation between years in fine-scale spatial genetic substructuring of populations. Our results indicate that the viability of Phlox pilosa can be best ensured by protecting and creating populations of at least 1000–2000 flowering ramets.     

Reduced pollinator service and elevated pollen limitation at the geographic range limit of an annual plant

Mutualisms are well known to influence individual fitness and the population dynamics of partner species, but little is known about whether they influence species distributions and the location of geographic range limits. Here, we examine the contribution of plant-pollinator interactions to the geographic range limit of the California endemic plant Clarkia xantiana ssp. xantiana. We show that pollinator availability declined from the center to the margin of the geographic range consistently across four years of study. This decline in pollinator availability was caused to a greater extent by variation in the abundance of generalist rather than specialist bee pollinators. Climate data suggest that patterns of precipitation in the current and previous year drove variation in bee abundance because of its effects on cues for bee emergence in the current year and the abundance of floral resources in the previous year. Experimental floral manipulations showed that marginal populations had greater outcross pollen limitation of reproduction, in parallel with the decline in pollinator abundance. Although plants are self-compatible, we found no evidence that autonomous selfing contributes to reproduction, and thus no evidence that it alleviates outcross pollen limitation in marginal populations. Furthermore, we found no association between the distance to the range edge and selfing rate, as estimated from sequence and microsatellite variation, indicating that the mating system has not evolved in response to the pollination environment at the range periphery. Overall, our results suggest that dependence on pollinators for reproduction may be an important constraint limiting range expansion in this system.     

The demographic consequences of the cost of reproduction in ungulates

The cost of reproduction can generate covariation between demographic rates that can potentially influence demography and population dynamics in long-lived iteroparous species. However, there has been relatively little work linking the survival cost of reproduction and population dynamics. The apparent scarcity of information on this important link is potentially due to covariation between vital rates, which can substantially influence fluctuations in population size. In this paper we examine the opportunity for survival costs of reproduction to leave a dynamic signature using a simulation model based broadly on an ungulate life history. We find that an increase in the cost delays the onset of reproduction and reduces reproductive rates of young, but not of prime-age, females. Accordingly, the number of offspring produced declines and the interval between reproductive events increases among young females experiencing high cost. These effects are translated to an age structure skewed toward young ages and reduced population density. These results suggest that, by delaying reproduction when conditions deteriorate, females protect their survival during the critical first three years of life, after which the negative effect of reproduction on survival declines. Unless conditions for reproduction are severe, it is not profitable to delay reproduction beyond age 3 years due to the high risk of death before having a chance to reproduce. We also demonstrate that lack of adjustment of reproductive strategies to elevated levels of the cost of reproduction, for example due to rapid changes in environmental conditions, results in lower average density and longevity compared to females that have sufficient time to adjust to changes in the cost. This suggests that even moderate costs of reproduction may have a major negative effect on population dynamics of ungulates.     

A comparative seasonal study of two populations of Hypnea musciformis from the East and West Coasts of Florida,USA II. Photosynthetic and respiratory rates

Respiration rates of Hypnea musciformis (Wulfen) Lamouroux in Florida, USA, generally increased with increased temperature. Gulf coast H. musciformis respired at significantly higher rates than the Atlantic coast population, which exhibited a region of temperature independence between 24°–32°C. Respiration rates were highest in the fall and winter, during the periods of rapid growth. Respiration rates were lowest in the summer indicating a period of storage and low metabolism. Photosynthetic responses to various levels of light and temperature indicated that the Gulf coast population was more tolerant to high light intensities than the Atlantic coast population. Maximum photosynthetic responses for both populations occurred between 24° and 32°C which corresponds to the shallow slope region of the respiration-temperature curves. The results indicate that water temperature rather than light intensity is a significant factor in modifying seasonal photosynthetic capacities. The greatest seasonal variation in photosynthetic responses occurred at the light-temperature levels of highest responses while little seasonal variation was demonstrated at tolerance limits.     

Effects of Agricultural, Industrial, and Recreational Expansion on Frequency of Wildlife Law Violations in the Central Rocky Mountains, USA

Abstract: As human populations expand into previously unoccupied habitats, conflicts with wild vertebrate species are inevitable. On the basis of literature concerning human attitudes toward nature, we hypothesized that the intensity of conflicts and impacts would vary with the type of land use expansion. We tested our hypothesis by examining data on socioeconomic profiles of discrete expanding human populations in relation to the frequency of wildlife law violations in the central Rocky Mountains of the United States. We estimated possible human-related effects by comparing rates of wildlife law violations among populations varying in growth rates and socioeconomic status over an eleven-year period 1969–1980. We found that agrarian-based population centers had fewer violations per capita than areas with industrial-based (petroleum development) boom towns. Recreation-oriented (ski) boom towns had the lowest number of violations during periods of the most rapid growth. While the relationship between wildlife law violations and actual impact on animal population sizes remains unknown, our findings underscore the necessity of a priori recognition and planning by government and private agencies to combat potential harmful effects when expanding human population centers are likely to be characterized by immigrants with a low regard for wild species.     

Integrating quality and quantity of mutualistic service to contrast ant species protecting Ferocactus wislizeni

Generalized, facultative mutualisms are often characterized by great variation in the benefits provided by different partner species. This variation may be due to differences among species in the quality and quantity of their interactions, as well as their phenology. Many plant species produce extrafloral nectar, a carbohydrate-rich resource, to attract ant species that can act as "bodyguards" against a plant's natural enemies. Here, we explore differences in the quality and quantity of protective service that ants can provide a plant by contrasting the four most common ant visitors to Ferocactus wislizeni, an extrafloral nectary-bearing cactus in southern Arizona. The four species differ in abundance when tending plants, and in the frequency at which they visit plants. By adding surrogate herbivores (Manduca sexta caterpillars) to plants, we demonstrate that all four species recruit to and attack potential herbivores. However, their per capita effectiveness in deterring herbivores (measured as the inverse of the number of workers needed to remove half of the experimentally added caterpillars) differs. Using these among-species differences in quality (per capita effectiveness) and quantity (number of workers that visit a plant and frequency of visitation), we accurately predicted the variation in fruit production among plants with different histories of ant tending. We found that plant benefits (herbivore removal and maturation of buds and fruits) typically saturated at high levels of ant protection, although plants could be "well defended" via different combinations of interaction frequency, numbers of ant workers per interaction, and per capita effects. Our study documents variation among prospective mutualists, distinguishes the components of this variation, and integrates these components into a predictive measure of protection benefit to the plant. The method we used to average saturating benefits over time could prove useful for quantifying overall service in other mutualisms.     

Leaf traits are good predictors of plant performance across 53 rain forest species

We compared the leaf traits and plant performance of 53 co-occurring tree species in a semi-evergreen tropical moist forest community. The species differed in all leaf traits analyzed: leaf life span varied 11-fold among species, specific leaf area 5-fold, mass-based nitrogen 3-fold, mass-based assimilation rate 13-fold, mass-based respiration rate 15-fold, stomatal conductance 8-fold, and photosynthetic water use efficiency 4-fold. Photosynthetic traits were strongly coordinated, and specific leaf area predicted mass-based rates of assimilation and respiration; leaf life span predicted many other leaf characteristics. Leaf traits were closely associated with growth, survival, and light requirement of the species. Leaf investment strategies varied on a continuum trading off short-term carbon gain against long-term leaf persistence that, in turn, is linked to variation in whole-plant growth and survival. Leaf traits were good predictors of plant performance, both in gaps and in the forest understory. High growth in gaps is promoted by cheap, short-lived, and physiologically active leaves. High survival in the forest understory is enhanced by the formation of long-lived well protected leaves that reduce biomass loss by herbivory, mechanical disturbance, or leaf turnover. Leaf traits underlay this growth-survival trade-off; species with short-lived, physiologically active leaves have high growth but low survival. This continuum in leaf traits, through its effect on plant performance, in turn gives rise to a continuum in species' light requirements.