Photosynthetic resource-use efficiency and demographic variability in desert winter annual plants

We studied a guild of desert winter annual plants that differ in long-term variation in per capita reproductive success (lb, the product of per capita survival from germination to reproduction, l, times per capita reproduction of survivors, b) to relate individual function to population and community dynamics. We hypothesized that variation in lb should be related to species' positions along a trade-off between relative growth rate (RGR) and photosynthetic water-use efficiency (WUE) because lb is a species-specific function of growing-season precipitation. We found that demographically variable species have greater RGR and greater leaf carbon isotope discrimination (Delta, a proxy inversely related to WUE). We examined leaf nitrogen and photosynthetic characteristics and found that, in this system, variation in Delta is a function of photosynthetic demand rather than stomatal regulation of water loss. The physiological characteristics that result in low Delta in some species may confer greater photosynthetic performance during the reliably moist but low temperature periods that immediately follow winter rainfall in the Sonoran Desert or alternatively during cool periods of the day or early growing season. Conversely, while species with high Delta and high RGR exhibit low leaf N, they have high biomass allocation to canopy leaf area display. Such trait associations may allow for greater performance during the infrequent conditions where high soil moisture persists into warmer conditions, resulting in high demographic variance. Alternatively, high variance could arise from specialization to warm periods of the day or season. Population dynamic buffering via stress tolerance (low RGR and Delta) correlates negatively with buffering via seed banks, as predicted by bet-hedging theory. By merging analyses of population dynamics with functional trait relationships, we develop a deeper understanding of the physiological, ecological, and evolutionary mechanisms involved in population and community dynamics.     

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.     

Seasonal timing of first rain storms affects rare plant population dynamics

A major challenge in forecasting the ecological consequences of climate change is understanding the relative importance of changes to mean conditions vs. changes to discrete climatic events, such as storms, frosts, or droughts. Here we show that the first major storm of the growing season strongly influences the population dynamics of three rare and endangered annual plant species in a coastal California (USA) ecosystem. In a field experiment we used moisture barriers and water addition to manipulate the timing and temperature associated with first major rains of the season. The three focal species showed two- to fivefold variation in per capita population growth rates between the different storm treatments, comparable to variation found in a prior experiment imposing eightfold differences in season-long precipitation. Variation in germination was a major demographic driver of how two of three species responded to the first rains. For one of these species, the timing of the storm was the most critical determinant of its germination, while the other showed enhanced germination with colder storm temperatures. The role of temperature was further supported by laboratory trials showing enhanced germination in cooler treatments. Our work suggests that, because of species-specific cues for demographic transitions such as germination, changes to discrete climate events may be as, if not more, important than changes to season-long variables.     

Effects of life history and reproduction on recruitment time lags in reintroductions of rare plants

Reintroductions are important components of conservation and recovery programs for rare plant species, but their long-term success rates are poorly understood. Previous reviews of plant reintroductions focused on short-term (e.g., ≤3 years) survival and flowering of founder individuals rather than on benchmarks of intergenerational persistence, such as seedling recruitment. However, short-term metrics may obscure outcomes because the unique demographic properties of reintroductions, including small size and unstable stage structure, could create lags in population growth. We used time-to-event analysis on a database of unusually well-monitored and long-term (4–28 years) reintroductions of 27 rare plant species to test whether life-history traits and population characteristics of reintroductions create time-lagged responses in seedling recruitment (i.e., recruitment time lags [RTLs]), an important benchmark of success and indicator of persistence in reintroduced populations. Recruitment time lags were highly variable among reintroductions, ranging from <1 to 17 years after installation. Recruitment patterns matched predictions from life-history theory with short-lived species (fast species) exhibiting consistently shorter and less variable RTLs than long-lived species (slow species). Long RTLs occurred in long-lived herbs, especially in grasslands, whereas short RTLs occurred in short-lived subtropical woody plants and annual herbs. Across plant life histories, as reproductive adult abundance increased, RTLs decreased. Highly variable RTLs were observed in species with multiple reintroduction events, suggesting local processes are just as important as life-history strategy in determining reintroduction outcomes. Time lags in restoration outcomes highlight the need to scale success benchmarks in reintroduction monitoring programs with plant life-history strategies and the unique demographic properties of restored populations. Drawing conclusions on the long-term success of plant reintroduction programs is premature given that demographic processes in species with slow life-histories take decades to unfold.     

Double Allee Effects and Extinction in the Island Fox

Abstract:  An Allee effect (AE) occurs in populations when individuals suffer a decrease in fitness at low densities. If a fitness component is reduced (component AE), per capita population growth rates may decline as a consequence (demographic AE) and extinction risk is increased. The island fox ( Urocyon littoralis ) is endemic to six of the eight California Channel Islands. Population crashes have coincided with an increase in predation by Golden Eagles ( Aquila chrysaetos ). We propose that AEs could render fox populations more sensitive and may be a likely explanation for their sharp decline. We analyzed demographic data collected between 1988 and 2000 to test whether fox density (1) influences survival and reproductive rates; (2) interacts with eagle presence and affects fox fitness parameters; and (3) influences per capita fox population trends. A double component AE simultaneously influenced survival (of adults and pups) and proportion of breeding adult females. The adult survival AE was driven by predation by eagles. These component AEs led to a demographic AE. Multiple-component AEs, a predation-driven AE, and the simultaneous occurrence of both component and demographic AEs in a mammal are all previously unreported processes. Populations below 7 foxes/km² could have suboptimal population growth rates due to the demographic AE, and AEs may have contributed to the dramatic declines in three fox populations. Because fox densities in critically endangered populations are well below this level, removing Golden Eagles appears necessary to prevent a predation-driven AE. Conservationists should also be aware of AEs when planning the release of captive foxes. More generally, our findings highlight the danger of overlooking AEs in the conservation of populations of rare or threatened species.     

Benefits of large broods by higher chick survival and better territories in a precocial shorebird

When reproductive success is constant in one breeding phase, different tactics that increase variation in reproductive success among individuals may evolve in other phases. For instance, in shorebirds, which usually have a limited clutch size of four eggs, variation in reproductive tactics among individuals is expected either before egg-laying (e.g. diverse mating systems) or after hatching of the young (e.g. diverse parental care). In this paper, I studied the pied avocet (Recurvirostra avosetta), a shorebird with a modal clutch size of four eggs, to test whether post-hatch chick adoption as an alternative tactic can be linked to increased variation in annual reproductive success. When predation was high, naturally adopting pairs produced more filial fledglings than did pairs not adopting chicks and not losing chicks to adoption. The number of filial fledglings increased with the number of adopted young, possibly through diluting the chances of predation on filial young. Experimental chick addition did not lead to more fledged young due to low brood integrity as shown by the frequent loss of chicks from some experimental broods. When predation was low, larger broods occupied feeding territories with higher prey abundance than smaller broods, possibly due to their dominance over smaller ones. Pairs that lost chicks to adoption (donors) fledged as many filial young in their broods as did non-adopters/non-donors, whereas the total number of donors’ filial fledglings, including those raised in adopting broods, approached that of adopters. These findings show, for the first time, that post-hatch alternative reproductive tactics can lead to variation in annual reproductive success and to higher success for some pairs even in species where past adaptations limit variation in reproductive success in a certain phase of reproduction.     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

Competitive females are successful females; phenotype,mechanism, and selection in a common songbird

In a variety of taxa, male reproductive success is positively related to the expression of costly traits such as large body size, ornaments, armaments, and aggression. These traits are thought to improve male competitive ability and, thus, access to limited reproductive resources. Females of many species also express competitive traits. However, we know very little about the consequences of individual variation in competitive traits and the mechanisms that regulate their expression in females. Consequently, it is currently unclear whether females express competitive traits owing to direct selection or as an indirect result of selection on males. Here, we examine females of a mildly dimorphic songbird (Junco hyemalis) to determine whether females show positive covariance in traits (morphology and behavior) that may be important in a competition. We also examine whether trait expression relates either to testosterone (T) in terms of mechanism or to reproductive success in terms of function. We found that larger females were more aggressive and that greater ability to produce T in response to a physiological challenge consisting of a standardized injection of gonadotropin-releasing hormone predicted some measures of female body size and aggression. Finally, we found that aggressive females had greater reproductive success. We conclude that T may influence female phenotype and that females may benefit from expressing a competitive phenotype. We also suggest that the mild dimorphism observed in many species may be due in part to direct selection on females rather than simply a correlated response to selection in males.     

Phenotypic correlates and survival consequences of male mating success in lek-mating greater prairie-chickens (<Emphasis Type="Italic">Tympanuchus cupido</Emphasis>)

Female choice and male–male aggression are two modes of sexual selection that can lead to elaboration of male morphological and behavioral traits. In lek-mating species, male mating success is often strongly skewed, and it is puzzling why variation in male traits is still observed given directional female choice. If male traits correlated with reproductive success are honest signals of male quality, there may be survival costs associated with the expression of those traits. In this study, we examined whether morphological, behavioral, and territorial traits are correlated with male mating success and survival in the lek-mating greater prairie-chicken (Tympanuchus cupido). We introduce a novel application of multinomial discrete choice models for analysis of female mate choice behavior. We found that behavioral and territorial attributes showed 6.5 times more variability among males than morphological traits. Both display and aggressive behaviors were strong predictors of male mating success, suggesting that both female choice and male–male aggression were important in determining mating success among male greater prairie-chickens. Moreover, annual survival of male prairie-chickens was independent of mating success and male traits. Females appear to be choosing males based on behavioral traits where large variation exists between males (coefficient of variation >30%). Behavioral traits were the most important factor in determining mating success of male prairie-chickens, but the mechanism underlying this relationship is unknown. In the future, experimental manipulations of male hormones or parasite loads could bridge the proximate mechanisms and ultimate consequences of factors mediating male mating success in lek-mating grouse.     

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.     

Linking fruit traits to variation in predispersal vertebrate seed predation, insect seed predation, and pathogen attack

The importance of vertebrates, invertebrates, and pathogens for plant communities has long been recognized, but their absolute and relative importance in early recruitment of multiple coexisting tropical plant species has not been quantified. Further, little is known about the relationship of fruit traits to seed mortality due to natural enemies in tropical plants. To investigate the influences of vertebrates, invertebrates, and pathogens on reproduction of seven canopy plant species varying in fruit traits, we quantified reductions in fruit development and seed germination due to vertebrates, invertebrates, and fungal pathogens through experimental removal of these enemies using canopy exclosures, insecticide, and fungicide, respectively. We also measured morphological fruit traits hypothesized to mediate interactions of plants with natural enemies of seeds. Vertebrates, invertebrates, and fungi differentially affected predispersal seed mortality depending on the plant species. Fruit morphology explained some variation among species; species with larger fruit and less physical protection surrounding seeds exhibited greater negative effects of fungi on fruit development and germination and experienced reduced seed survival integrated over fruit development and germination in response to vertebrates. Within species, variation in seed size also contributed to variation in natural enemy effects on seed viability. Further, seedling growth was higher for seeds that developed in vertebrate exclosures for Anacardium excelsum and under the fungicide treatment for Castilla elastica, suggesting that predispersal effects of natural enemies may carry through to the seedling stage. This is the first experimental test of the relative effects of vertebrates, invertebrates, and pathogens on seed survival in the canopy. This study motivates further investigation to determine the generality of our results for plant communities. If there is strong variation in natural enemy attack among species related to differences in fruit morphology, then quantification of fruit traits will aid in predicting the outcomes of interactions between plants and their natural enemies. This is particularly important in tropical forests, where high species diversity makes it logistically impossible to study every plant life history stage of every species.     

Potential Misuse of Avian Density as a Conservation Metric

Abstract: Effective conservation metrics are needed to evaluate the success of management in a rapidly changing world. Reproductive rates and densities of breeding birds (as a surrogate for reproductive rate) have been used to indicate the quality of avian breeding habitat, but the underlying assumptions of these metrics rarely have been examined. When birds are attracted to breeding areas in part by the presence of conspecifics and when breeding in groups influences predation rates, the effectiveness of density and reproductive rate as indicators of habitat quality is reduced. It is beneficial to clearly distinguish between individual‐ and population‐level processes when evaluating habitat quality. We use the term reproductive rate to refer to both levels and further distinguish among levels by using the terms per capita fecundity (number of female offspring per female per year, individual level) and population growth rate (the product of density and per capita fecundity, population level). We predicted how density and reproductive rate interact over time under density‐independent and density‐dependent scenarios, assuming the ideal free distribution model of how birds settle in breeding habitats. We predicted population density of small populations would be correlated positively with both per capita fecundity and population growth rate due to the Allee effect. For populations in the density‐dependent growth phase, we predicted no relation between density and per capita fecundity (because individuals in all patches will equilibrate to the same success rate) and a positive relation between density and population growth rate. Several ecological theories collectively suggest that positive correlations between density and per capita fecundity would be difficult to detect. We constructed a decision tree to guide interpretation of positive, neutral, nonlinear, and negative relations between density and reproductive rates at individual and population levels.     

Variation in signal timing behavior: implications for male attractiveness and sexual selection

Mate choice often takes place in group settings, such as leks or choruses, in which numerous individuals display and compete for mates simultaneously. In addition to well-known preferences for male traits like size and signaling rate, females of group-displaying species often show preferences that are based on the relative timing of male signals, generally preferring the leading signal. Variation in male signal timing behavior may therefore affect male attractiveness and, ultimately, reproductive success. I used acoustic communication in green treefrogs (Hyla cinerea) to assess the amount of signal timing variation found in natural choruses and to test hypotheses about the sources of variation in signal timing behavior. I recorded dyads of vocally interacting males in the field to describe patterns of variation in signal timing behavior. Incorporating information about female signal timing preferences, I also assessed the amount of variation in signal timing that is likely to be selectively neutral, as well as the amount that is likely under selection by female choice because it involves attractive or unattractive signal placement. I show that there is considerable variation in signal timing behavior, particularly involving neutral signal timing placement. I also show that between-male variation in other traits (size, signal period) is partly linked to variation in signal timing behavior and discuss these findings in terms of male attractiveness and sexual selection in group-displaying organisms.     

Living at the Edge: Local versus Positional Factors in the Long‐Term Population Dynamics of an Endangered Orchid

Abstract: Populations at the margin of geographic ranges of distribution have been considered more vulnerable than central ones, but recent reviews have caste doubt on this generalization. We examined the reproductive and demographic performance of a rare Euroasiatic orchid (Cypripedium calceolus) at its southwesterly range limit and compared our findings with those of previous studies of nine central populations at the center of the orchid's range. We sought to test the central‐marginal model and to evaluate factors involved in long‐term performance of forest Eurosiberian species with peripheral populations in southern European mountains. We characterized (structure, temporal fluctuations, herbivory, reproductive success, and recruitment at different habitats) four Pyrenean populations of C. calceolus of different sizes (5–3500 ramets) and monitored three of them for up to 13 years. Two quantitative stochastic models (count data and matrix models) were used to assess population trends and viability and the effect of herbivory. Contrary to expectations, and despite the negative effect of sporadic events of herbivory, the peripheral populations we studied (except the smallest one) performed similarly or better than populations occurring in central part of the species’ range in terms of reproductive success and population growth rates. Landscape changes over the last 50 years suggest that natural reforestation could be involved in the success of this plant at its southern limit. Forest expansion in the mountain regions of southern Europe may provide new opportunities for plants with geographic distributions centered mainly at higher latitudes and give some hope for their recovery in future scenarios dominated by biodiversity loss.     

Sexual selection and reproductive careers in mandrills (Mandrillus sphinx)

In sexually dimorphic, polygynous species, where males provide little parental care and competition between males for access to fertile females is high, sexual selection theory predicts sex differences in age-specific reproductive output and mortality profiles, and greater variance in lifetime reproductive success in males than in females. We examined age-specific reproductive output, mortality patterns and the extent and causes of variation in reproductive success for a semi-free-ranging colony of mandrills (Mandrillus sphinx, Cercopithecidae) in Franceville, Gabon, using long-term (20 year) demographic records and microsatellite parentage analysis. Although differences in the demography and feeding ecology of this closed, provisioned colony, in comparison with wild mandrills, limit interpretation of our results, sex differences in reproductive output and mortality showed the patterns predicted by sexual selection theory. Mortality was higher in males than in females after sexual maturity, and lifespan was significantly shorter in males (mean 14 year) than in females (>22 year). Age at first reproduction was significantly earlier in females (mean 4.2 year) than in males (11.6 year), and male reproductive output declined earlier. All females of breeding age produced offspring; while only 17 of 53 sexually mature males (32%) sired. Males sired a maximum of 41 offspring, versus 17 in females, and variance in male reproductive output was significantly greater than in females at all ages. The most important influence on variation in lifetime reproductive output in both sexes was joint variation between length of the breeding period and reproductive rate, due to lower reproductive rates in younger animals. Finally, social rank significantly influenced reproductive output in both sexes: high-ranking females began their reproductive careers earlier and had a higher subsequent reproductive rate than low-ranking females, while males that achieved top rank during their career sired far more offspring than males that did not.Electronic Supplementary Material Supplementary material is available for this article at     

Causes and consequences of unequal seedling production in forest trees: a case study in red oaks

Inequality in reproductive success has important implications for ecological and evolutionary dynamics, but lifetime reproductive success is challenging to measure in long-lived species such as forest trees. While seed production is often used as a proxy for overall reproductive success, high mortality of seeds and the potential for trade-offs between seed number and quality draw this assumption into question. Parentage analyses of established seedlings can bring us one step closer to understanding the causes and consequences of variation in reproductive success. In this paper we demonstrate a new method for estimating individual seedling production and average percentage germination, using data from two mixed-species populations of red oaks (Quercus rubra, Q. velutina, Q. falcata, and Q. coccinea). We use these estimates to examine the distribution of female reproductive success and to test the relationship between seedling number and individual seed production, age, and growth rate. We show that both seed and seedling production are highly skewed, roughly conforming to zero-inflated lognormal distributions, rather than to the Poisson or negative-binomial distributions often assumed by population genetics analyses. While the number of established offspring is positively associated with mean annual seed production, a lower proportion of seeds from highly fecund individuals become seedlings. Our red oak populations also show evidence of trade-offs between growth rate and reproductive success. The high degree of inequality in seedling production shown here for red oaks, and by previous studies in other species, suggests that many trees may be more vulnerable to genetic drift than previously thought, if immigration in limited by fragmentation or other environmental changes.     

Are functional traits good predictors of demographic rates? Evidence from five neotropical forests

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.     

Fourth-corner correlation is a score test statistic in a log-linear trait–environment model that is useful in permutation testing

Ecologists wish to understand the role of traits of species in determining where each species occurs in the environment. For this, they wish to detect associations between species traits and environmental variables from three data tables, species count data from sites with associated environmental data and species trait data from data bases. These three tables leave a missing part, the fourth-corner. The fourth-corner correlations between quantitative traits and environmental variables, heuristically proposed 20 years ago, fill this corner. Generalized linear (mixed) models have been proposed more recently as a model-based alternative. This paper shows that the squared fourth-corner correlation times the total count is precisely the score test statistic for testing the linear-by-linear interaction in a Poisson log-linear model that also contains species and sites as main effects. For multiple traits and environmental variables, the score test statistic is proportional to the total inertia of a doubly constrained correspondence analysis. When the count data are over-dispersed compared to the Poisson or when there are other deviations from the model such as unobserved traits or environmental variables that interact with the observed ones, the score test statistic does not have the usual chi-square distribution. For these types of deviations, row- and column-based permutation methods (and their sequential combination) are proposed to control the type I error without undue loss of power (unless no deviation is present), as illustrated in a small simulation study. The issues for valid statistical testing are illustrated using the well-known Dutch Dune Meadow data set.     

Reproductive seasonality is a poor predictor of receptive synchrony and male reproductive skew among nonhuman primates

Among nonhuman primates, male reproductive skew (i.e., the distribution of reproductive success across males) appears to be affected primarily by receptive synchrony and the number of males per group. These factors have been assumed to depend on reproductive seasonality, with strong seasonality increasing receptive synchrony, which in turn reduces the strength of male monopolization associated with more males and lower skew. Here we tested the importance of reproductive seasonality for 26 populations representing 15 species living in multimale groups. We obtained data from the literature on paternity, number of males per group, receptive synchrony, and three measures of seasonality of reproduction. We analyzed these data using bivariate regressions and hierarchical regression by sets and controlled for the effect of evolutionary relationships using phylogenetic generalized least squares. As expected, alpha male paternity decreased as the number of males per group increased as well as with increasing female receptive synchrony. Reproductive seasonality did not explain variation in reproductive skew over and above the variation explained by synchrony and the number of males. Reproductive seasonality alone only explained a small proportion of the variation in skew, and there was no strong association between reproductive seasonality and synchrony. The effects of receptive synchrony and reproductive seasonality as well as their link were reduced if we excluded captive populations. These results indicate that across primates male reproductive skew is related to the number of competitors in a group and that seasonality does not reliably predict synchrony or male reproductive skew.     

Individual and environmental determinants of reproductive success in male tree swallow (Tachycineta bicolor)

Evaluating the contribution of individual and environmental determinants of reproductive success is essential to improve our understanding of sexual selection. In socially monogamous bird species with high rates of extrapair paternity, traits or environmental contexts affecting the number of within-pair young (WPY) produced by males can differ from those affecting the number of extrapair young fathered (EPY). Here, we use a 4-year dataset collected in contrasted environments to assess the factors affecting male reproductive success in tree swallows (Tachycineta bicolor), a species showing high levels of extrapair paternity. Our analyses revealed that the number of WPY was higher under better environmental conditions, while the number of EPY was mainly related to male characteristics. Males nesting in more intensive agricultural areas had fewer WPY produced and a lower reproductive success. Also, males breeding earlier in the season had more WPY. The presence of parasites reduced males’ reproductive success, mainly by reducing the number of EPY. The influence of male phenotype varied according to population density: Tarsus length variation had a greater effect on reproductive success at low population density than at high density, while wing length was also positively related to the number of EPY, more so at high than at low density. Altogether, our results suggest a complex interplay between individual and environmental determinants of reproductive success and imply that sexual selection dynamics varies depending on environmental contexts.