Showing papers in "The American Naturalist in 2002"

Phylogenetic analysis and comparative data: a test and review of evidence

Abstract: The question is often raised whether it is statistically necessary to control for phylogenetic associations in comparative studies. To investigate this question, we explore the use of a measure of phylogenetic correlation, lambda, introduced by Pagel (1999), that normally varies between 0 (phylogenetic independence) and 1 (species' traits covary in direct proportion to their shared evolutionary history). Simulations show lambda to be a statistically powerful index for measuring whether data exhibit phylogenetic dependence or not and whether it has low rates of Type I error. Moreover, lambda is robust to incomplete phylogenetic information, which demonstrates that even partial information on phylogeny will improve the accuracy of phylogenetic analyses. To assess whether traits generally show phylogenetic associations, we present a quantitative review of 26 published phylogenetic comparative data sets. The data sets include 103 traits and were chosen from the ecological literature in which debate about the need for phylogenetic correction has been most acute. Eighty-eight percent of data sets contained at least one character that displayed significant phylogenetic dependence, and 60% of characters overall (pooled across studies) showed significant evidence of phylogenetic association. In 16% of tests, phylogenetic correlation could be neither supported nor rejected. However, most of these equivocal results were found in small phylogenies and probably reflect a lack of power. We suggest that the parameter lambda be routinely estimated when analyzing comparative data, since it can also be used simultaneously to adjust the phylogenetic correction in a manner that is optimal for the data set, and we present an example of how this may be done.

Evolutionary origins and ecological consequences of endophyte symbiosis with grasses.

Abstract: Over the past 20 yr much has been learned about a unique symbiotic interaction between fungal endophytes and grasses. The fungi (Clavicipitaceae, Ascomycota) grow intercellularly and systemically in aboveground plant parts. Vertically transmitted asexual endophytes forming asymptomatic infections of cool‐season grasses have been repeatedly derived from sexual species that abort host inflorescences. The phylogenetic distribution of seed‐transmitted endophytes is strongly suggestive of cocladogenesis with their hosts. Molecular evidence indicates that many seed‐transmitted endophytes are interspecific hybrids. Superinfection may result in hyphal fusion and parasexual recombination. Most endophytes produce one or more alkaloid classes that likely play some role in defending the host plant against pests. Hybridization may have led to the proliferation of alkaloid‐production genes among asexual endophytes, favoring hybrids. The ergot alkaloid ergovaline, lolitrems, and lolines are produced by only a ...

Immune defense and host life history.

Abstract: Recent interest has focused on immune response in an evolutionary context, with particular attention to disease resistance as a life-history trait, subject to trade-offs against other traits such as reproductive effort. Immune defense has several characteristics that complicate this approach, however; for example, because of the risk of autoimmunity, optimal immune defense is not necessarily maximum immune defense. Two important types of cost associated with immunity in the context of life history are resource costs, those related to the allocation of essential but limited resources, such as energy or nutrients, and option costs, those paid not in the currency of resources but in functional or structural components of the organism. Resource and option costs are likely to apply to different aspects of resistance. Recent investigations into possible trade-offs between reproductive effort, particularly sexual displays, and immunity have suggested interesting functional links between the two. Although all organisms balance the costs of immune defense against the requirements of reproduction, this balance works out differently for males than it does for females, creating sex differences in immune response that in turn are related to ecological factors such as the mating system. We conclude that immune response is indeed costly and that future work would do well to include invertebrates, which have sometimes been neglected in studies of the ecology of immune defense.

Speciation by natural and sexual selection: models and experiments.

Abstract: A large number of mathematical models have been developed that show how natural and sexual selection can cause prezygotic isolation to evolve. This article attempts to unify this literature by identifying five major elements that determine the outcome of speciation caused by selection: a form of disruptive selection, a form of isolating mechanism (assortment or a mating preference), a way to transmit the force of disruptive selection to the isolating mechanism (direct selection or indirect selection), a genetic basis for increased isolation (a one- or two-allele mechanism), and an initial condition (high or low initial divergence). We show that the geographical context of speciation (allopatry vs. sympatry) can be viewed as a form of assortative mating. These five elements appear to operate largely independently of each other and can be used to make generalizations about when speciation is most likely to happen. This provides a framework for interpreting results from laboratory experiments, which are found to agree generally with theoretical predictions about conditions that are favorable to the evolution of prezygotic isolation.

Self-organization of vegetation in arid ecosystems

Abstract: Scientists are still searching for possible unifying mechanisms to explain this range of spatial patterns (Tongway and Ludwig 2001), and an important question of this research is whether this range is the result of preexisting environmental heterogeneity, the result of spatial selforganization, or both (Klausmeier 1999; Couteron and Lejeune 2001; HilleRisLambers et al. 2001; Von Hardenberg et al. 2001). Here, we contribute to the ongoing debate about vegetation pattern formation in arid ecosystems by presenting novel, spatially explicit model analyses and results, extending on the work of HilleRisLambers et al. (2001). Our results show that these different vegetation patterns observed in arid ecosystems might all be the result of spatial self-organization, caused by one single mechanism: water infiltrates faster into vegetated ground than into bare soil, leading to net displacement of surface water to vegetated patches. This model differs from earlier model results (Klausmeier 1999; Couteron and Lejeune 2001; HilleRisLambers et al. 2001; Von Hardenberg et al. 2001) primarily in two ways: it is fully mechanistic, and it treats the lateral flow of water above and below the soil as separate, not independent, variables. Although the current model greatly simplifies the biophysics of arid systems, it can reproduce the whole range of distinctive vegetation patterns as observed in arid ecosystems, indicating that the proposed mechanism might be generally applicable. We further show that self-organized vegetation patterns can persist far into regions of high aridity, where plants would become extinct if homogeneously distributed, pointing to the importance of this mechanism for maintaining productivity of arid ecosystems (Noy-Meir 1973). Our analyses are based on the model first developed in HilleRisLambers et al. (2001)

Metapopulation Structure Favors Plasticity over Local Adaptation

Abstract: We describe a model for the evolutionary consequences of plasticity in an environmentally heterogeneous metapopulation in which specialists for each of two alternative environments and one plastic type are initially present. The model is similar to that proposed by Moran (1992) but extends her work to two sites. We show that with migration between sites the plastic type is favored over local specialists across a broad range of parameter space. The plastic type may dominate or be fixed even in an environmentally uniform site, and even if the plasticity has imperfect accuracy or bears some cost such that a local specialist has higher fitness in that site, as long as there is some migration between sites with different distributions of environmental states. These results suggest that differences among taxa in dispersal and hence realized migration rates may play a heretofore unrecognized role in their patterns of adaptive population differentiation. Migration relaxes the thresholds for both environmental heterogeneity and accuracy of plastic response above which plasticity is favored. Furthermore, small changes in response accuracy can dramatically and abruptly alter the evolutionary outcome in the metapopulation. A fitness cost to plasticity will substantially reduce the range of conditions in which the plastic type will prevail only if the cost is both large and global rather than environment specific.

Comparing classical community models: theoretical consequences for patterns of diversity.

Abstract: Mechanisms proposed to explain the maintenance of species diversity within ecological communities of sessile organisms include niche differentiation mediated by competitive trade-offs, frequency dependence resulting from species-specific pests, re- cruitment limitation due to local dispersal, and a speciation- extinction dynamic equilibrium mediated by stochasticity (drift). While each of these processes, and more, have been shown to act in particular communities, much remains to be learned about their relative importance in shaping community-level patterns. We used a spatially-explicit, individual-based model to assess the effects of each of these processes on species richness, relative abundance, and spatial patterns such as the species-area curve. Our model com- munities had an order-of-magnitude more individuals than any previous such study, and we also developed a finite-size scaling analysis to infer the large-scale properties of these systems in order to establish the generality of our conclusions across system sizes. As expected, each mechanism can promote diversity. We found some qualitative differences in community patterns across com- munities in which different combinations of these mechanisms operate. Species-area curves follow a power law with short-range dispersal and a logarithmic law with global dispersal. Relative- abundance distributions are more even for systems with compet- itive differences and trade-offs than for those in which all species are competitively equivalent, and they are most even when fre- quency dependence (even if weak) is present. Overall, however, communities in which different processes operated showed sur- prisingly similar patterns, which suggests that the form of com- munity-level patterns cannot in general be used to distinguish among mechanisms maintaining diversity there. Nevertheless, pa- rameterization of models such as these from field data on the

Toward a causal explanation of plant invasiveness: seedling growth and life-history strategies of 29 pine (Pinus) species.

Abstract: We studied 29 pine (Pinus) species to test the hypothesis that invasive species in disturbed habitats have distinct attributes. Seedling relative growth rate (RGR) and measures of invasiveness were positively associated across species as well as within phylogenetically independent contrasts. High RGR, small seed masses, and short generation times characterize pine species that are successful invaders in disturbed habitats. Discriminant analysis and logistic regression revealed that RGR was the most significant factor among these life-history traits separating invasive and noninvasive species. We also explored the causes of differences in RGR among invasive and noninvasive species. While net assimilation rate, leaf mass ratio, and specific leaf area (SLA) were all found to be contributing positively to RGR, SLA was found to be the main component responsible for differences in RGR between invasive and noninvasive pines. We investigated differences in SLA further by studying leaf anatomy, leaf density, and leaf thickness. We also evaluated relative leaf production rate as an important aspect of SLA. We proposed a hypothetical causal network of all relevant variables.

Species Invasions Exceed Extinctions on Islands Worldwide: A Comparative Study of Plants and Birds

Abstract: Species richness is decreasing at a global scale. At subglobal scales, that is, within any defined area less extensive than the globe, species richness will increase when the number of nonnative species becoming naturalized is greater than the number of native species becoming extinct. Determining whether this has occurred is usually difficult because detailed records of species extinctions and naturalizations are rare; these records often exist, however, for oceanic islands. Here we show that species richness on oceanic islands has remained relatively unchanged for land birds, with the number of naturalizations being roughly equal to the number of extinctions, and has increased dramatically for vascular plants, with the number of naturalizations greatly exceeding the number of extinctions. In fact, for plants, the net number of species on islands has approximately doubled. We show further that these patterns are robust to differences in the history of human occupation of these islands and to th...

Why alien invaders succeed: support for the escape-from-enemy hypothesis.

Abstract: Successful biological invaders often exhibit enhanced performance following introduction to a new region. The traditional explanation for this phenomenon is that natural enemies (e.g., competitors, pathogens, and predators) present in the native range are absent from the introduced range. The purpose of this study was to test the escape-from-enemy hypothesis using the perennial plant Silene latifolia as a model system. This European native was introduced to North America in the 1800s and subsequently spread to a large part of the continent. It is now considered a problematic weed of disturbed habitats and agricultural fields in the United States and Canada. Surveys of 86 populations in the United States and Europe revealed greater levels of attack by generalist enemies (aphids, snails, floral herbivores) in Europe compared with North America. Two specialists (seed predator, anther smut fungus) that had dramatic effects on plant fitness in Europe were either absent or in very low frequency in North America. Overall, plants were 17 times more likely to be damaged in Europe than in North America. Thus, S. latifolia's successful North American invasion can, at least in part, be explained by escape from specialist enemies and lower levels of damage following introduction.

Species Richness and Altitude: A Comparison between Null Models and Interpolated Plant Species Richness along the Himalayan Altitudinal Gradient, Nepal

Abstract: We compare different null models for species richness patterns in the Nepalese Himalayas, the largest altitudinal gradient in the world. Species richness is estimated by interpolation of presences between the extreme recorded altitudinal ranges. The number of species in 100-m altitudinal bands increases steeply with altitude until 1,500 m above sea level. Between 1,500 and 2,500 m, little change in the number of species is observed, but above this altitude, a decrease in species richness is evident. We simulate different null models to investigate the effect of hard boundaries and an assumed linear relationship between species richness and altitude. We also stimulate the effect of interpolation when incomplete sampling is assumed. Some modifications on earlier simulations are presented. We demonstrate that all three factors in combination may explain the observed pattern in species richness. Estimating species richness by interpolating species presence between maximum and minimum altitudes creates an artificially steep decrease in species richness toward the ends of the gradient. The addition of hard boundaries and an underlying linear trend in species richness is needed to simulate the observed broad pattern in species richness along altitude in the Nepalese Himalayas.

Costs of phenotypic plasticity.

Abstract: Phenotypically plastic organisms display alternative phenotypes in different environments. It is widely appreciated that possessing alternative phenotypes can affect fitness. However, some investigators have suggested that simply carrying the ability to be plastic could also affect fitness. Evolutionary models suggest that high costs of plasticity could constrain the evolution of optimal phenotypes. However, costs (and limits) of plasticity are primarily hypothetical. Little empirical evidence exists to show that increased plasticity leads to reduced growth and development, leads to increased developmental instability, or limits the ability of organisms to produce more extreme phenotypes. I used half‐sib families of larval wood frogs (Rana sylvatica) reared in outdoor mesocosms to examine how tadpoles altered behavioral, morphological, and life‐historical traits in response to larval dragonfly predators (Anax longipes). The predators induced lower activity and the development of relatively large...

Nitrogen in Insects: Implications for Trophic Complexity and Species Diversification

Abstract: Disparities in nutrient content (nitrogen and phosphorus) between herbivores and their plant resources have lately proven to have major consequences for herbivore success, consumer‐driven nutrient cycling, and the fate of primary production in ecosystems. Here we extend these findings by examining patterns of nutrient content between animals at higher trophic levels, specifically between insect herbivores and predators. Using a recently compiled database on insect nutrient content, we found that predators exhibit on average 15% greater nitrogen content than herbivores. This difference persists after accounting for variation from phylogeny and allometry. Among herbivorous insects, we also found evidence that recently derived lineages (e.g., herbivorous Diptera and Lepidoptera) have, on a relative basis, 15%–25% less body nitrogen than more ancient herbivore lineages (e.g., herbivorous Orthoptera and Hemiptera). We elaborate several testable hypotheses for the origin of differences in nitrogen con...

Coexistence in metacommunities: the regional similarity hypothesis.

Abstract: Species richness has historically been studied with a separation between smalland large-scale processes. Species diversity has been approached, on the one hand, from a local perspective, based on niche theory (Pianka 1966; MacArthur and Levins 1967; Schoener 1974), and on the other hand, from a regional perspective, through island biogeography (MacArthur and Wilson 1967), with no strong interactions between these two levels. At the local scale, interactions between competing species constrain diversity, and coexistence is a function of niche dimensions and resource heterogeneity (MacArthur and Levins 1967) or differences in species life-history traits as in colonization-competition trade-off models (Hastings 1980; Tilman 1994). At the regional scale, the theory of island biogeography (MacArthur and Wilson 1967) ignores local dynamics and considers local diversity as the result of regional processes such as chance events of immigration and extinction. There are no limits to diversity except those arising from the size of the regional species pool (continent size) and the constraints on immigration events (continent-island distance). This apparent contradiction has been named “MacArthur’s paradox” (Schoener 1983; Loreau and Mouquet 1999) because MacArthur’s contribution has been central in both niche theory (Mac-

Individual Covariation in Life‐History Traits: Seeing the Trees Despite the Forest

Abstract: We investigated the influence of age on survival and breeding rates in a long‐lived species Rissa tridactyla using models with individual random effects permitting variation and covariation in fitness components among individuals. Differences in survival or breeding probabilities among individuals are substantial, and there was positive covariation between survival and breeding probability; birds that were more likely to survive were also more likely to breed, given that they survived. The pattern of age‐related variation in these rates detected at the individual level differed from that observed at the population level. Our results provided confirmation of what has been suggested by other investigators: within‐cohort phenotypic selection can mask senescence. Although this phenomenon has been extensively studied in humans and captive animals, conclusive evidence of the discrepancy between population‐level and individual‐level patterns of age‐related variation in life‐history traits is extremely ...

Developmental thresholds and the evolution of reaction norms for age and size at life-history transitions.

Abstract: It is quite common in studies of life‐history plasticity to find a negative relationship between the age at which various life‐history transitions occur and the growth conditions under which individuals develop In particular, high growth typically results in earlier transitions, often at a larger size Here, we use a relatively general optimization model for age and size at life‐history transitions to argue that current life‐history theory cannot adequately explain these results Specifically, most such theory requires key assumptions that are unlikely to be generally met This suggests that some important component of the biology of many organisms must be missing from many of the models in life‐history theory We suggest that this missing component might be the phenomenon of developmental thresholds There are at least two different types of developmental thresholds possible, and we incorporate these into our general optimality model to demonstrate how they can cause a negative relationship be

Patchy Reaction‐Diffusion and Population Abundance: The Relative Importance of Habitat Amount and Arrangement

Abstract: A discrete reaction‐diffusion model was used to estimate long‐term equilibrium populations of a hypothetical species inhabiting patchy landscapes to examine the relative importance of habitat amount and arrangement in explaining population size. When examined over a broad range of habitat amounts and arrangements, population size was largely determined by a pure amount effect (proportion of habitat in the landscape accounted for >96% of the total variation compared to <1% for the arrangement main effect). However, population response deviated from a pure amount effect as coverage was reduced below 30%–50%. That deviation coincided with a persistence threshold as indicated by a rapid decline in the probability of landscapes supporting viable populations. When we partitioned experimental landscapes into sets of “above” and “below” persistence threshold, habitat arrangement became an important factor in explaining population size below threshold conditions. Regression analysis on below‐threshold la...

Ecological specialization and susceptibility to disturbance: conjectures and refutations.

Abstract: Niche breadth of species has been hypothesized to be associated with species' responses to disturbance. Disturbance is usually believed to affect specialists negatively, while generalists are believed to benefit from disturbance; we call this the "specialization-disturbance" hypothesis. We also propose an associated hypothesis (the "specialization-asymmetry-disturbance" hypothesis) under which both specialization and asymmetry of interactions would explain species' responses to disturbance. We test these hypotheses using data from a plant-pollinator system that has been grazed by cattle (i.e., a biological disturbance) in southern Argentina. We quantified specialization in species interactions, specialization of interaction partners, and species' responses to disturbance. We found no relationship between degree of specialization and a species' response to disturbance. We also found that plant-pollinator interactions tend to be asymmetric in this system; there was no relationship between the degree of specialization of a given species and the degree of specialization of its interaction partners. However, asymmetry of interactions did not explain the variability in species' responses to disturbance. Thus, both hypotheses are rejected by our data. Possible reasons include failure to assess crucial resources, substantial direct effects of disturbance, inaccurate measures of specialization, difficulty detecting highly nonlinear relationships, and limitations of a nonexperimental approach. Or, in fact, there may be no relationship between specialization and response to disturbance.

Multiple receivers, multiple ornaments, and a trade-off between agonistic and epigamic signaling in a widowbird.

Abstract: Sexual displays often involve several different ornamental traits. Yet most indicator models of sexual selection based on a single receiver (usually a choosy female) find that multiple handicap signals should be unstable.Here we study reasons for this contradiction, analyzing signal function, signal content, and trade-offs between signals in the polygynous red-collared widowbird Euplectes ardens. Males have both a long, graduated tail and a red carotenoid collar badge. Territory-holding "residents" have slightly shorter tails than the nonbreeding "floaters," but their carotenoid collars are 40% larger, and they have (on the basis of reflectance spectrometry and objective colorimetry) a 23-nm more long-wave ("redder") hue than floaters. This corroborates experimental evidence that the red collar is selected by male contest competition, whereas female choice is based almost exclusively on male tail length. Tail length is negatively correlated with the carotenoid signal, which together with body size and condition explains 55% of the variation in tail length. The trade-off in tail length and carotenoid investment is steeper among residents, suggesting an interaction with costs of territory defense. We propose that the "multiple receiver hypothesis" can explain the coexistence of multiple handicap signals. Furthermore, the trade-off between signal expressions might contribute to the inverse relation between nuptial tail elongation and coloration in the genus Euplectes (bishops and widowbirds).

Population Dynamics and Mutualism: Functional Responses of Benefits and Costs

Abstract: We develop an approach for studying population dynamics resulting from mutualism by employing functional responses based on density-dependent benefits and costs. These functional responses express how the population growth rate of a mutualist is modified by the density of its partner. We present several possible dependencies of gross benefits and costs, and hence net effects, to a mutualist as functions of the density of its partner. Net effects to mutualists are likely a monotonically saturating or unimodal function of the density of their partner. We show that fundamental differences in the growth, limitation, and dynamics of a population can occur when net effects to that population change linearly, unimodally, or in a saturating fashion. We use the mutualism between senita cactus and its pollinating seed-eating moth as an example to show the influence of different benefit and cost functional responses on population dynamics and stability of mutualisms. We investigated two mechanisms that may alter this mutualism's functional responses: distribution of eggs among flowers and fruit abortion. Differences in how benefits and costs vary with density can alter the stability of this mutualism. In particular, fruit abortion may allow for a stable equilibrium where none could otherwise exist.

A Comparative Test of the Adaptive Plasticity Hypothesis: Relationships between Habitat and Phenotype in Anuran Larvae

Abstract: The hypothesis that phenotypic plasticity is maintained by divergent natural selection acting across different environments predicts that populations and species exposed to highly variable environments will express high levels of plasticity. I tested this prediction by measuring the behavioral and morphological responses to aeshnid dragonfly larvae of 16 tadpole species and asking whether predator‐induced plasticity is greater in species that experience more variable densities of predators in nature. Tadpole phenotypes were measured in a series of similar experiments in outdoor artificial ponds carried out over a 9‐yr period. I quantified tadpole habitats by soliciting evaluations by seven to 36 experienced field observers for each species. There were large differences among species in phenotype, mostly in agreement with earlier descriptions. Nearly all species responded to dragonflies by decreasing activity and body length relative to overall body size and by increasing relative tail fin depth,...

Delayed dispersal as a route to breeding: Territorial inheritance, safe havens, and ecological constraints

Abstract: The relative roles of ecological constraints, the benefits of philopatry, and the role of life history continue to be debated in the evolution of natal philopatry and cooperative breeding. We compare three routes to breeding: departing to search for territories as a floater, staying and queuing to inherit the natal territory, or queuing and eventually shifting to a neighboring vacancy. Our model assumed a dominance-structured population. It quantifies the benefits of philopatry for varying-rank subordinates and contrasts it against the benefit of dispersal. We apply the model to data on Siberian jay Perisoreus infaustus, a species in which retained offspring do not help at the nest. The results indicate that territorial inheritance plays a small role in this species (presumably due to inbreeding avoidance), and territory acquisition is less constrained for dispersing than philopatric offspring. Nevertheless, small family groups-one or, at the most, two same-sex queuers-are predicted to form because philopatric offspring gain nepotistic benefits that improve their survival. This fits with data on group sizes and supports the idea of the natal territory as a safe haven for waiting for breeding opportunities. We also discuss our predictions in the light of ecological constraints and clarify recent confusingly different predictions on the role of habitat saturation as an explanation for delayed dispersal and cooperative breeding. We argue that "ecological constraint" is too wide a term to yield useful predictive power and that it is more appropriate to examine the consequences of specific life-history traits on the success of dispersers.

How Do Sinking Phytoplankton Species Manage to Persist

Abstract: Phytoplankton require light for photosynthesis. Yet, most phytoplankton species are heavier than water and therefore sink. How can these sinking species persist? Somehow, the answer should lie in the turbulent motion that redisperses sinking phytoplankton over the vertical water column. Here, we show, using a reaction-advection-diffusion equation of light-limited phytoplankton, that there is a turbulence window sustaining sinking phytoplankton species in deep waters. If turbulent diffusion is too high, phytoplankton are mixed to great depths, and the depth-averaged light conditions are too low to allow net positive population growth. Conversely, if turbulent diffusion is too low, sinking phytoplankton populations end up at the ocean floor and succumb in the dark. At intermediate levels of turbulent diffusion, however, phytoplankton populations can outgrow both mixing rates and sinking rates. In this way, the reproducing population as a whole can maintain a position in the well-lit zone near the top of the water column, even if all individuals within the population have a tendency to sink. This theory unites earlier classic results by Sverdrup and Riley as well as our own recent findings and provides a new conceptual framework for the understanding of phytoplankton dynamics under the influence of mixing processes.

Dispersal: risk spreading versus local adaptation.

Abstract: I investigate how risk spreading in stochastic environments and adaptation to permanent properties of local habitats interplay in the simultaneous evolution of dispersal and habitat specialization. In a simple two‐patch model, I find many types of locally evolutionarily stable attractors of dispersal and of a trait involved in habitat specialization, including a single habitat specialist and a coalition of two specialists with low dispersal, a generalist with high dispersal, and several types of dispersal polymorphisms. In general, only one attractor is a global evolutionarily stable strategy (ESS). In addition to the ESS analysis, I also present some examples of the dynamics of evolution that exhibit adaptive diversification by evolutionary branching.

Sex chromosomes and sexual selection in poeciliid fishes.

Abstract: We propose that the evolution of female preferences can be strongly influenced by linkage of attractive male traits to the Y chromosome and female preferences to the X chromosome in male heterogametic species. Such linkage patterns are predicted by models of the evolution of sexually antagonistic genes. Subsequent recombination of attractive male characters from the Y to the X would create physical linkage between attractive male trait and preference. A literature survey shows that Y linkage of potentially sexually antagonistic traits is common in poeciliid fishes and other species with sex chromosomes that are not well differentiated, but may also occur in taxa with degenerate Y chromosomes. In the guppy, attractive male traits are primarily Y and X linked; a literature review of the inheritance of sex‐limited attractive male characters suggests that 16 are Y linked, 24 recombine between the X and Y, two are X linked, and two are autosomal. Crosses and backcrosses between high female preference...

Testing for Environmentally Induced Bias in Phenotypic Estimates of Natural Selection: Theory and Practice

Abstract: Measuring natural selection has been a fundamental goal of evolutionary biology for more than a century, and techniques developed in the last 20 yr have provided relatively simple means for biologists to do so. Many of these techniques, however, share a common limitation: when applied to phenotypic data, environmentally induced covariances between traits and fitness can lead to biased estimates of selection and misleading predictions about evolutionary change. Utilizing estimates of breeding values instead of phenotypic data with these methods can eliminate environmentally induced bias, although this approach is more difficult to implement. Despite this potential limitation to phenotypic methods and the availability of a potential solution, little empirical evidence exists on the extent of environmentally induced bias in phenotypic estimates of selection. In this article, we present a method for detecting bias in phenotypic estimates of selection and demonstrate its use with three independent da...

On the vegetative biomass partitioning of seed plant leaves, stems, and roots.

Abstract: A central goal of comparative life‐history theory is to derive the general rules governing growth, metabolic allocation, and biomass partitioning. Here, we use allometric theory to predict the relationships among annual leaf, stem, and root growth rates (GL, GS, and GR, respectively) across a broad spectrum of seed plant species. Our model predicts isometric scaling relationships among all three organ growth rates: \documentclass{aastex} \usepackage{amsbsy} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{bm} \usepackage{mathrsfs} \usepackage{pifont} \usepackage{stmaryrd} \usepackage{textcomp} \usepackage{portland,xspace} \usepackage{amsmath,amsxtra} \usepackage[OT2,OT1]{fontenc} ewcommand\cyr{ \renewcommand\rmdefault{wncyr} \renewcommand\sfdefault{wncyss} \renewcommand\encodingdefault{OT2} ormalfont \selectfont} \DeclareTextFontCommand{\textcyr}{\cyr} \pagestyle{empty} \DeclareMathSizes{10}{9}{7}{6} \begin{document} \landscape $G_{\mathrm{L}\,}\propto G_{\mathrm{S}\,}\propto G_{\mathr...

Species richness, environmental correlates, and spatial scale: a test using South African birds

Abstract: Energy and habitat heterogeneity are important correlates of spatial variation in species richness, though few investigations have sought to determine simultaneously their relative influences. Here we use the South African avifauna to examine the extent to which species richness is related to these variables and how these relationships depend on spatial grain. Taking spatial autocorrelation and area effects into account, we find that primary productivity, precipitation, absolute minimum temperature, and, at coarser resolutions, habitat heterogeneity account for most of the variation in species richness. Species richness and productivity are positively related, whereas the relationship between potential evapotranspiration (PET) and richness is unimodal. This is largely because of the constraining effects of low rainfall on productivity in high-PET areas. The increase in the importance of vegetation heterogeneity as an explanatory variable is caused largely by an increase in the range of vegetation heterogeneity included at coarse resolutions and is probably also a result of the positive effects of environmental heterogeneity on species richness. Our findings indicate that species richness is correlated with, and hence likely a function of, several variables, that spatial resolution and extent must be taken into account during investigations of these relationships, and that surrogate measures for productivity should be interpreted cautiously.

Coexistence and relative abundance in annual plant assemblages: the roles of competition and colonization.

Abstract: Although an interspecific trade-off between competitive and colonizing ability can permit multispecies coexistence, whether this mechanism controls the structure of natural systems remains unresolved. We used models to evaluate the hypothesized importance of this trade-off for explaining coexistence and relative abundance patterns in annual plant assemblages. In a nonspatial model, empirically derived competition-colonization trade-offs related to seed mass were insufficient to generate coexistence. This was unchanged by spatial structure or interspecific variation in the fraction of seeds dispersing globally. These results differ from those of the more generalized competition-colonization models because the latter assume completely asymmetric competition, an assumption that appears unrealistic considering existing data for annual systems. When, for heuristic purposes, completely asymmetric competition was incorporated into our models, unlimited coexistence was possible. However, in the resulting abundance patterns, the best competitors/poorest colonizers were the most abundant, the opposite of that observed in natural systems. By contrast, these natural patterns were produced by competition-colonization models where environmental heterogeneity permitted species coexistence. Thus, despite the failure of the simple competition-colonization trade-off to explain coexistence in annual plant systems, this trade-off may be essential to explaining relative abundance patterns when other processes permit coexistence.