Wintering hummingbirds in Alabama and Florida: species diversity, sex and age ratios, and site fidelity

ABSTRACT Over the past several decades, there have been numerous reports of hummingbirds wintering in the southeastern United States. However, little is known about the species present and their relative abundance. From November 1998 to March 2008, we examined the species diversity, sex and age ratios, and site fidelity of hummingbirds wintering in southern Alabama and northern Florida. We captured and banded 1598 individuals representing 10 species, and the most frequently captured species were Rufous Hummingbirds (Selasphorus rufus; 51.6%), Ruby‐throated Hummingbirds (Archilochus colubris; 23.5%), and Black‐chinned Hummingbirds (Archilochus alexandri; 16.9%). Other species captured included Buff‐bellied Hummingbirds (Amazilia yucatanensis), Calliope Hummingbirds (Stellula calliope), Allen's Hummingbirds (Selasphorus sasin), Broad‐tailed Hummingbirds (Selasphorus platycercus), Broad‐billed Hummingbirds (Cynanthus latirostris), Anna's Hummingbirds (Calypte anna), and Costa's Hummingbirds (Calypte costae). Most hummingbirds (71.8%) were captured in December and January. For most species, sex ratios were male‐biased for juveniles and female‐biased for adults, indicating possible differential mortality. Of 1598 hummingbirds captured, 144 representing five species returned to the same wintering location at least once. Female Rufous Hummingbirds (20.4% of individuals captured) exhibited the greatest site fidelity. Recaptures of banded Rufous Hummingbirds in autumn and early winter revealed that some individuals moved south into Alabama or Florida from Tennessee, northern Georgia, and northern Louisiana. Same‐season recaptures of banded Rufous Hummingbirds suggest that their spring migration route is west along the Gulf Coast. Our results suggest that Alabama and Florida are viable overwintering areas for several species of hummingbirds, with numbers of species and individuals higher than previously recognized. However, more study is needed to confirm migration routes and to determine if Ruby‐throated Hummingbirds wintering in our study area are year‐round residents or migrants.     

An estimator of the Opportunity for Selection that is independent of mean fitness

Variation among individuals in number of offspring (fitness, k) sets an upper limit to the evolutionary response to selection. This constraint is quantified by Crow's Opportunity for Selection (I), which is the variance in relative fitness (I = σ²_k/(u_k)²). Crow's I has been widely used but remains controversial because it depends on mean offspring number in a sample (). Here, I used a generalized Wright-Fisher model that allows for unequal probabilities of producing offspring to evaluate behavior of Crow's I and related indices under a wide range of sampling scenarios. Analytical and numerical results are congruent and show that rescaling the sample variance (s²_k) to its expected value at a fixed removes dependence of I on mean offspring number, but the result still depends on choice of . A new index is introduced, Δ_I = Π– E(Î_drift) = Π– 1/, which makes Î independent of sample without the need for variance rescaling. Δ_I has a straightforward interpretation as the component of variance in relative fitness that exceeds that expected under a null model of random reproductive success. Δ_I can be used to directly compare estimates of the Opportunity for Selection for samples from different studies, different sexes, and different life stages.     

Quantitative genetics of temperature performance curves of Neurospora crassa

Earth's temperature is increasing due to anthropogenic CO emissions; and organisms need either to adapt to higher temperatures, migrate into colder areas, or face extinction. Temperature affects nearly all aspects of an organism's physiology via its influence on metabolic rate and protein structure, therefore genetic adaptation to increased temperature may be much harder to achieve compared to other abiotic stresses. There is still much to be learned about the evolutionary potential for adaptation to higher temperatures, therefore we studied the quantitative genetics of growth rates in different temperatures that make up the thermal performance curve of the fungal model system Neurospora crassa. We studied the amount of genetic variation for thermal performance curves and examined possible genetic constraints by estimating the G -matrix. We observed a substantial amount of genetic variation for growth in different temperatures, and most genetic variation was for performance curve elevation. Contrary to common theoretical assumptions, we did not find strong evidence for genetic trade-offs for growth between hotter and colder temperatures. We also simulated short-term evolution of thermal performance curves of N. crassa, and suggest that they can have versatile responses to selection.     

The genetic architecture of maternal effects across ontogeny in the red deer

Maternal effects, either environmental or genetic in origin, are an underappreciated source of phenotypic variance in natural populations. Maternal genetic effects have the potential to constrain or enhance the evolution of offspring traits depending on their magnitude and their genetic correlation with direct genetic effects. We estimated the maternal effect variance and its genetic component for 12 traits expressed over the life history in a pedigreed population of wild red deer (morphology, survival/longevity, breeding success). We only found support for maternal genetic effect variance in the two neonatal morphological traits: birth weight ( = 0.31) and birth leg length ( = 0.17). For these two traits, the genetic correlation between maternal and direct additive effects was not significantly different from zero, indicating no constraint to evolution from genetic architecture. In contrast, variance in maternal genetic effects enhanced the additive genetic variance available to respond to natural selection. Maternal effect variance was negligible for late-life traits. We found no evidence for sex differences in either the direct or maternal genetic architecture of offspring traits. Our results suggest that maternal genetic effect variance declines over the lifetime, but also that this additional heritable genetic variation may facilitate evolutionary responses of early-life traits.     

EXTINCTION-RECOLONIZATION DYNAMICS IN THE MYCOPHAGOUS BEETLE PHALACRUS SUBSTRIATUS

The population structure of the mycophagous beetle Phalacrus substriatus is characterized by many small, local populations interconnected by migration over a small spatial scale (10 × 75 m²). Each local P. substriatus population has a relatively short expected persistence time, but persistence of the species occurs due to a balance between frequent local extinctions and recolonizations. This nonequilibrium population structure can have profound effects on how the genetic variation is structured between and within populations. Theoretical models have stated that the genetic differentiation among local populations will be enhanced relative to an island model at equilibrium if the number of colonizers is less than approximately twice the number of migrants among local populations. To study these effects, a set of 50 local P. substriatus populations were surveyed over a four-year period to record any naturally occurring extinctions and recolonizations. The per population colonization and extinction rate were 0.237 and 0.275, respectively. Mark-recapture techniques were used to estimate a number of demographic parameters: local population size (N = 11.1), migration rate , number of colonizers (k = 4.0), and the probability of common origin of colonizers (φ = 0.5). The theoretically predicted level of differentiation among local populations (measured as Wright's F_ST) was 0.070. Genetic data obtained from an electrophoretic survey of seven polymorphic loci gave an estimated degree of differentiation of 0.077. There was thus a good agreement between the empirical results and the theoretical predictions. Young populations had significantly higher levels of differentiation than old, more established populations . The extinction-recolonization dynamics resulted in an overall increase in the genetic differentiation among local populations by c. 40%. The global effective population size was also reduced by c. 55%. The results give clear evidence to how nonequilibrium processes shape the genetic structure of populations.     

Genetic Characteristics of Restored Elk Populations in Kentucky

Translocations are a common management practice to restore or augment populations. Understanding the genetic consequences of translocation efforts is important for the long-term health of restored populations. The restoration of elk (Cervus canadensis) to Kentucky, USA, included source stocks from 6 western states, which were released at 8 sites in southeastern Kentucky during 1997–2002. We assessed genetic diversity in restored herds and compared genetic similarity to source stocks based on 15 microsatellite DNA loci. Genetic variation in the restored populations was comparable to source stocks ( allelic richness = 3.52 and 3.50; expected heterozygosity = 0.665 and 0.661 for restored and source, respectively). Genetic differentiation among all source and restored populations ranged from 0.000 to 0.065 for pairwise F_ST and 0.034 to 0.161 for pairwise Nei's D_A. Pairwise genetic differentiation and Bayesian clustering revealed that stocks from Utah and North Dakota, USA, contributed most to restored populations. Other western stocks appeared less successful and were not detected with our data, though our sampling was not exhaustive. We also inferred natural movements of elk among release sites by the presence of multiple genetic stocks. The success of the elk restoration effort in Kentucky may be due, in part, to the large number of elk (n = 1,548), repeated releases, and use of diverse source stocks. Future restoration efforts for elk in the eastern United States should consider the use of multiple stock sources and a large number of individuals. In addition, preservation of genetic samples of founder stock will enable detailed monitoring in the future. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

ADAPTIVE RADIATION AND GENETIC DIFFERENTIATION IN THE HAWAIIAN SILVERSWORD ALLIANCE (COMPOSITAE: MADIINAE)

The Hawaiian silversword alliance consists of the three genera Dubautia, Argyroxiphium, and Wilkesia, and is a classic example of adaptive radiation in an insular setting. Genetic variation and interspecific genetic differentiation based on ten enzyme loci are described for Dubautia and Wilkesia. Genetic identities among species span the range of values expected from interpopulation comparisons within a single species (I = 0.90–1.00) to those typical of interspecific comparisons . Genetic-identity values correspond to biogeographic distribution and morphological distinctiveness, supporting a correlation of increasing genetic distance associated with the time of separation among lineages. It may be inferred that the high genetic identities observed within the Hawaiian Madiinae and other island plant groups are due to limited time spans available for taxa to accumulate new genetic variation through mutation. It appears that species may remain genetically similar (I > 0.90) even after time spans on the order of magnitude of 1,000,000 years.     

Resource use,energetic profitability,and behavioral decisions in migrant rufous hummingbirds

Summary Migrant Rufous Hummingbirds (Selasphorus rufus) defend nectar resources at stopover sites while replenishing fat reserves needed for migratory flights. During late summer in the Sandia Mountains, central New Mexico, they defend the wasp- and bee-pollinated Scrophularia montana from other hummingbirds. Both hummingbirds and hymenopterans exploit Scrophularia nectar during the early part of its flowering period. As summer colony growth increases the densities of the eusocial hymenopterans by 100–150%, their exploitation of Scrophularia nectar lowers its mean standing crop in flowers by 200–300%. Sometime during the summer, Rufous Hummingbirds abandon and do not further use this resource for the remaining 3–4 weeks of its flowering period. The abandonment always occurs when the mean standing crop of nectar is approximately 0.2–0.3 L/ flower. This paper describes a model of Rufous Hummingbird energetics, that shows abandonment occurred 1–3 days after they passed the threshold at which the resource could have provided their minimum daily energy requirements. I suggest that constraints imposed by a highly competitive social environment severely reduced the options available to the hummingbirds, and caused them to continue to defend a resource that could no longer meet their energetic requirements.     

SMALL POPULATION GENETIC VARIABILITY AT LOCI UNDER STABILIZING SELECTION

Genetic variability at a locus under stabilizing selection in a finite population is investigated using analytic methods and computer simulations. Three measures are examined: the number of alleles k, heterozygosity H, and additive genetic variance Vg. A nearly-neutral theory results. The composite parameter S = NV_M/V_s (where N is the population size, V_M the variance of new mutant allelic effects and V_s the weakness of stabilizing selection) figures prominently in the results. The equilibrium heterozygosity is similar to that of strictly neutral theory, H = 4N_μc/ (1 + 4N_μc), except that μ_c = where c is about 0.5. Simulations corroborate except for very low N. Genetic variability attains similar equilibrium values at both a “lone” locus and at an “embedded” locus. This agrees with my earlier work concerning molecular clock rates. These results modify the neutralist interpretation of data concerning genetic variability and genetic distances between populations. Low H values are proportional not to N but to . This may explain the narrow observed range of H among species. Heterozygosities need not be highly correlated to genetic variances. Genetic variances are not highly dependent on population size except in very small populations which are difficult to sample without bias because the smallest populations go extinct the fastest. Nearly neutral evolution will not be easily distinguished from strictly neutral theory under the Hudson-Kreitman-Aguade inter-/intraspecific variation ratio test, since a similar effective mutation rate holds for genetic distances and D = 2μ_ct, where . As with strictly neutral theory, comparisons across loci should show D and H to be positively correlated because of the shared μ_c. But unlike neutral theory, for a given locus, comparisons across species should show D and H to be negatively correlated. There is no obvious threshold of population size below which genetic variability inevitably declines. Extinction depends on both genetic variation and natural selection. Neither theory nor observation presently indicates the measure of genetic variability (k, H, V_G or other) that best indicates vulnerability of a small population to extinction.     

Technical note: Measures of differentiation for quantitative traits

Studies of anthropological genetics and bioarcheology often examine the degree of among-group variation in quantitative traits such as craniometrics and anthropometrics. One comparative index of among-group differentiation is the minimum value of Wright's $F_{\mathrm{ST}}$ as estimated from quantitative traits. This measure has been used in certain population-genetic applications such as comparison with $F_{\mathrm{ST}}$ estimated from genetic data, although some inferences are limited by how well the data and study design fit the underlying population-genetic model. In many cases, all that is needed is a simple measure of among-group variation. One such measure is $R^2$, the proportion of total phenotypic variation accounted for by among-group phenotypic variation, a measure easily obtained from analysis of variance and regression methods. This paper shows that $R^2$ and minimum $F_{\mathrm{ST}}$ are closely related as $\mathrm{Min}{F}_{\mathrm{ST}}\approx R^2/\left(2-R^2\right)$. $R^2$ is computationally easy and may be useful in cases where all we need is a simple measure of relative among-group differentiation.     

Phenotypic evolution in stochastic environments: The contribution of frequency- and density-dependent selection

Understanding how environmental variation affects phenotypic evolution requires models based on ecologically realistic assumptions that include variation in population size and specific mechanisms by which environmental fluctuations affect selection. Here we generalize quantitative genetic theory for environmentally induced stochastic selection to include general forms of frequency- and density-dependent selection. We show how the relevant fitness measure under stochastic selection relates to Fisher's fundamental theorem of natural selection, and present a general class of models in which density regulation acts through total use of resources rather than just population size. In this model, there is a constant adaptive topography for expected evolution, and the function maximized in the long run is the expected factor restricting population growth. This allows us to generalize several previous results and to explain why apparently “-selected” species with slow life histories often have low carrying capacities. Our joint analysis of density- and frequency-dependent selection reveals more clearly the relationship between population dynamics and phenotypic evolution, enabling a broader range of eco-evolutionary analyses of some of the most interesting problems in evolution in the face of environmental variation.     

THE MAINTENANCE OF POLYGENIC VARIATION IN FINITE POPULATIONS

Models of the maintenance of genetic variance in a polygenic trait have usually assumed that population size is infinite and that selection is weak. Consequently, they will overestimate the amount of variation maintained in finite populations. I derive approximations for the equilibrium genetic variance, in finite populations under weak stabilizing selection for triallelic loci and for an infinite “rare alleles” model. These are compared to results for neutral characters, to the “Gaussian allelic” model, and to Wright's approximation for a biallelic locus under arbitrary selection pressures. For a variety of parameter values, the three-allele, Gaussian, and Wrightian approximations all converge on the neutral model when population size is small. As expected, far less equilibrium genetic variance can be maintained if effective population size, N, is on the order of a few hundred than if N is infinite. All of the models predict that comparisons among populations with N less than about 10⁴ should show substantial differences in . While it is easier to maintain absolute when alleles interact to yield dominance or overdominance for fitness, less additivity also makes more susceptible to differences in N. I argue that experimental data do not seem to reflect the predicted degree of relationship between N and . This calls into question the ability of mutation-selection balance or simple balancing selection to explain observed . The dependence of on N could be used to test the adequacy of mutation-selection balance models.     

RISK OF POPULATION EXTINCTION FROM FIXATION OF NEW DELETERIOUS MUTATIONS

The fixation of new deleterious mutations is analyzed for a randomly mating population of constant size with no environmental or demographic stochasticity. Mildly deleterious mutations are far more important in causing loss of fitness and eventual extinction than are lethal and semilethal mutations in populations with effective sizes, N_e, larger than a few individuals. If all mildly deleterious mutations have the same selection coefficient, s against heterozygotes and 2s against homozygotes, the mean time to extinction, , is asymptotically proportional to for 4N_es > 1. Nearly neutral mutations pose the greatest risk of extinction for stable populations, because the magnitude of selection coefficient that minimizes is about ? = 0.4/N_e. The influence of variance in selection coefficients among mutations is analyzed assuming a gamma distribution of s, with mean and variance . The mean time to extinction increases with variance in selection coefficients if is near ?, but can decrease greatly if is much larger than ?. For a given coefficient of variation of , the mean time to extinction is asymptotically proportional to for . When s is exponentially distributed, (c = 1) is asymptotically proportional to . These results in conjunction with data on the rate and magnitude of mildly deleterious mutations in Drosophila melanogaster indicate that even moderately large populations, with effective sizes on the order of N_e = 10³, may incur a substantial risk of extinction from the fixation of new mutations.     

THE EFFECT OF INBREEDING IN DIPLOID AND TETRAPLOID POPULATIONS OF EPILOBIUM ANGUSTIFOLIUM (ONAGRACEAE): IMPLICATIONS FOR THE GENETIC BASIS OF INBREEDING DEPRESSION

The partial dominance model for the evolution of inbreeding depression predicts that tetraploids should exhibit less inbreeding depression than their diploid progenitors. We tested this prediction by comparing the magnitude of inbreeding depression in tetraploid and diploid populations of the herbaceous perennial Epilobium angustifolium (Onagraceae). Inbreeding depression was estimated in the greenhouse for three tetraploid and two diploid populations at four life stages. The mating system of a tetraploid population was estimated and compared to a previous estimate for diploids. Tetraploids showed less inbreeding depression than diploids at all life history stages, and these differences were significant for seed-set and cumulative fitness, but not for germination, survival, or plant dry mass at nine weeks. This result suggests that the genetic basis of inbreeding depression may differ among life stages. The primary selfing rate of the tetraploid population was r = 0.43, which is nearly identical to that of a diploid population (r = 0.45), indicating that differences in inbreeding depression between diploids and tetraploids are probably not due to differences in the mating system. Cumulative inbreeding depression, calculated from the four life history stages, was significantly higher for diploids () than for tetraploids (), supporting the partial dominance model of inbreeding depression.     

Effects of partial selfing on the equilibrium genetic variance,mutation load,and inbreeding depression under stabilizing selection

The mating system of a species is expected to have important effects on its genetic diversity. In this article, we explore the effects of partial selfing on the equilibrium genetic variance V_g, mutation load L, and inbreeding depression δ under stabilizing selection acting on a arbitrary number n of quantitative traits coded by biallelic loci with additive effects. When the ratio is low (where U is the total haploid mutation rate on selected traits) and effective recombination rates are sufficiently high, genetic associations between loci are negligible and the genetic variance, mutation load, and inbreeding depression are well predicted by approximations based on single‐locus models. For higher values of and/or lower effective recombination, moderate genetic associations generated by epistasis tend to increase V_g, L, and δ, this regime being well predicted by approximations including the effects of pairwise associations between loci. For yet higher values of and/or lower effective recombination, a different regime is reached under which the maintenance of coadapted gene complexes reduces V_g, L, and δ. Simulations indicate that the values of V_g, L, and δ are little affected by assumptions regarding the number of possible alleles per locus.     

THE EFFECT OF SOFT SELECTION ON THE VARIABILITY OF A QUANTITATIVE TRAIT

The equilibrium phenotypic variance of a normally distributed quantitative character P under soft selection is studied. This character is assumed to undergo Gaussian stabilizing selection W(p, x) = exp[–(p – x)²/2w²]. The environmentally determined optimum (x) is a normal variable with variance s². A stable equilibrium with is found, so that increases both with increasing environmental heterogeneity and with increasing local intensity of stabilizing selection. It is shown that both genetic and environmental components of the variance are selected until this equilibrium is reached. Habitat selection, supposed to be normal (with variance H²) around the optimum, also increases the value. Nevertheless, relatively intense local stabilizing selection (w < s) and accurate habitat choice (H < s) are required for the initial spread and the evolutionary stability of this habitat selection.     

The effects of hurricanes on the stochastic population growth of the endemic epiphytic orchid Broughtonia cubensis living in Cuba

We carried out a posthurricane evaluation of Broughtonia cubensis (Lindl.) Cogn., an endemic Cuban epiphytic orchid, after Hurricane Ivan (2004). We studied the transient responses in the stochastic dynamics of the species at three different sites over 13 successive years (2006–2019), monitored plot inventories (464 individuals in 10 transects) and built stochastic population models. The deterministic stochastic growth rate values () did not significantly differ (F = 2.76; p > 0.076) among the three sites over the 2006–2019 period. The long-term stochastic growth rate was 0.973 [0.932, 1.034]. The matrix elements that had the largest effect on were the transition to and stasis within the largest size class. Transient responses explained an average of 86% of the variation in the observed population growth rates , compared to 4% of the variation in the vital rates . Because transient dynamics are dependent on the population size composition, we ran extinction risk analyses under two scenarios: a population composed mainly of juveniles and another composed mainly of adults. There was little risk of falling below the quasi-extinction threshold before 25 year for both juveniles and adults. However, the risk of quasi-extinction was almost certain for both size classes by 80 year. We also simulated the effect of increasing the hurricane occurrence probability over 80 year on the population. There was little risk of extinction before 20 year in the baseline model, but there was a significant risk of extinction within 5 year when 90% of the individuals were affected by a new hurricane event.     

Effective size of density‐dependent populations in fluctuating environments

Reliable estimates of effective population size are of central importance in population genetics and evolutionary biology. For populations that fluctuate in size, harmonic mean population size is commonly used as a proxy for (multi‐) generational effective size. This assumes no effects of density dependence on the ratio between effective and actual population size, which limits its potential application. Here, we introduce density dependence on vital rates in a demographic model of variance effective size. We derive an expression for the ratio in a density‐regulated population in a fluctuating environment. We show by simulations that yearly genetic drift is accurately predicted by our model, and not proportional to as assumed by the harmonic mean model, where N is the total population size of mature individuals. We find a negative relationship between and N. For a given N, the ratio depends on variance in reproductive success and the degree of resource limitation acting on the population growth rate. Finally, our model indicate that environmental stochasticity may affect not only through fluctuations in N, but also for a given N at a given time. Our results show that estimates of effective population size must include effects of density dependence and environmental stochasticity.