Spatiotemporal relationship between adult census size and genetic population size across a wide population size gradient

Adult census population size (N) and effective number of breeders (N_b) are highly relevant for designing effective conservation strategies. Both parameters are often challenging to quantify, however, making it of interest to determine whether one parameter can be generalized from the other. Yet, the spatiotemporal relationship between N and N_b has not been well characterized empirically in many taxa. We analysed this relationship for 5–7 consecutive years in twelve brook trout populations varying greatly in N (49‐10032) and N_b (3‐567) and identified major environmental variables affecting the two parameters. N or habitat size alone explained 47–57% of the variance in N_b, and N_b was strongly correlated with effective population size. The ratio N_b/N ranged from 0.01 to 0.45 and increased at small N or following an annual decrease in N, suggesting density‐dependent constraints on N_b. We found no evidence for a consistent, directional difference between variability in N_b and/or N_b/N among small and large populations; however, small populations had more varying temporal variability in N_b/N ratios than large populations. Finally, N_b and N_b/N were 2.5‐ and 2.3‐fold more variable among populations than temporally within populations. Our results demonstrate a clear linkage between demographic and evolutionary parameters, suggesting that N_b could be used to approximate N (or vice versa) in natural populations. Nevertheless, using one variable to infer the other to monitor trends within populations is less recommended, perhaps even less so in small populations given their less predictable N_b vs. N dynamics.     

Comparative biogeography reveals differences in population genetic structure of five species of stream fishes

The distribution of genetic variation in Texas stream fishes has been shaped by a complex mix of historical and anthropogenic factors. Although Texas was not glaciated during the Pleistocene, the rise in sea level following this epoch isolated formerly connected drainages. More recently, the construction of dams, modifications of stream systems, and the release of commercially raised fish have influenced the patterns of genetic diversity. To examine how these different factors have impacted Texas stream fishes, we compared the genetic structure of five species of fish spanning two families and inhabiting two adjacent drainages: Lepomis megalotis, Lepomis cyanellus, Cyprinella lutrensis, Cyprinella venusta, and Campostoma anomalum. Our analyses of the mitochondrial D‐Loop show that genetic patterns differ strongly across species. A phylogeographical split between the Brazos and Trinity drainages was seen in conspecific populations of Lepomis species and is probably the result of the historical separation of these river systems. In contrast, contemporary ecological and anthropogenic factors, such as the desiccation of streams during summer, and the translocation of bait fish, appear to have a stronger influence on the genetic patterns in the remaining species. The contrasting results demonstrate the importance of using a multi‐species, comparative approach for landscape genetic studies as single species patterns may not be representative of others and thus may obscure differential effects of historical versus recent events as well as natural versus anthropogenic forces. By comparing closely related species that differ in their life history and economic importance it may be possible to disentangle the relative roles of historical, intrinsic, and anthropogenic influences on different organisms within a region. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Convergence, divergence, and the effect of congeners on body size ratios in stream fishes

Abstract Using collections from the years 1892–1999, I determined maximum standard length within each of 1030 populations of riffle-inhabiting darters ( Etheostoma spp.) representing five species from 788 sites in Illinois. Each site contained one to four riffle-inhabiting species of Etheostoma . Based on maximum-sized individuals in each collection, I calculated a ratio of standard lengths for all sympatric species pairs. Null models were developed using random pairings of body size measurements from sites with only one species of riffle Etheostoma to test whether body size ratios of sympatric species pairs are larger, indicating divergence, or smaller, indicating convergence, than ratios generated from allopatric populations. Results suggest that two of nine species pairs tend to converge in body size when sympatric. This suggests that convergence, in some cases, may facilitate the persistence of similar species in diverse communities. When variation in sympatric congener number is accounted for, the E. caeruleum : E. spectabile interaction results in significantly increasing size ratios as congener number increases. Etheostoma caeruleum and E. spectabile are the most ecologically similar and evolutionarily closely related species in the dataset. The divergence within this pair, associated with sympatric congener number, suggests that the degree of divergent character displacement may be indirectly enhanced by community structure.     

Concurrent habitat and life history influences on effective/census population size ratios in stream-dwelling trout

Lower effective sizes (N(e)) than census sizes (N) are routinely documented in natural populations, but knowledge of how multiple factors interact to lower N(e)/N ratios is often limited. We show how combined habitat and life-history influences drive a 2.4- to 6.1-fold difference in N(e)/N ratios between two pristine brook trout (Salvelinus fontinalis) populations occupying streams separated by only 750 m. Local habitat features, particularly drainage area and stream depth, govern trout biomass produced in each stream. They also generate higher trout densities in the shallower stream by favoring smaller body size and earlier age-at-maturity. The combination of higher densities and reduced breeding site availability in the shallower stream likely leads to more competition among breeding trout, which results in greater variance in individual reproductive success and a greater reduction in N(e) relative to N. A similar disparity between juvenile or adult densities and breeding habitat availability is reported for other species and hence may also result in divergent N(e)/N ratios elsewhere. These divergent N(e)/N ratios between adjacent populations are also an instructive reminder for species conservation programs that genetic and demographic parameters may differ dramatically within species.     

Morphological differences between two ecologically similar sympatric fishes

Morphological differentiation and microhabitat segregation of two ecologically similar populations of pelagic planktivorous fishes, Coregonus albula and the smaller Coregonus fontanae, were studied in Lake Stechlin (northern Germany). Both populations performed diel vertical migrations, although C. fontanae was always situated in deeper pelagic water than C. albula both during day and night. Landmark‐based geometric morphometrics revealed that sympatric C. albula and C. fontanae differ in external morphology, with main differences found in head length and eye position, as well as in length and width of the caudal peduncle. Moreover, while C. albula has a similar morphology over all sizes, the shape of C. fontanae changes with size. Accordingly, the morphology of the two is most different at smaller size. Although the morphological differences may reflect adaptations to the slightly differing microhabitats of the two populations, there is no conclusive evidence that this correspondence between ecology and morphology is the main mechanism behind the coexistence of the closely related coregonids in Lake Stechlin.     

Fine-scale genetic structure in a free-living ungulate population

The fine-scale genetic structure of wild animal populations has rarely been analysed, yet is potentially important as a confounding factor in quantitative genetic and allelic association studies, as well as having implications for population dynamics, inbreeding and kin selection. In this study, we examined the extent to which the three spatial subunits, or hefts, of the Village Bay population of Soay sheep (Ovis aries) on St Kilda, Scotland, are genetically structured using data from 20 microsatellite and protein loci. Allele frequencies differed significantly among three hefts in all the study years we considered (1987-2000 inclusive). Small but significantly positive F(ST) and negative F(IS) values were observed in most years, indicating that the hefts are genetically differentiated, and that within each heft there is more observed heterozygosity than would be expected if each were an isolated breeding population. Males showed less fidelity to their natal heft, and as a consequence higher levels of relatedness within hefts were observed among females than among males. There was a significant negative relationship between geographical proximity and relatedness in pairwise comparisons involving females, and on average pairs of females located within 50 m of each other were related at the equivalent level of second cousins. Structure is therefore largely driven by incomplete postnatal dispersal by females. Mating appears to be random with respect to the spatial-genetic substructure of the hefts, and therefore genetic structure does not contribute to the overall rate of inbreeding in the population. However, genetic substructure can lead to allelic associations and generate environmental effects within lineages that have the potential to confound heritability analyses and allelic association studies.     

Fine-scale population genetic structure in a fission-fusion society

Nonrandom patterns of mating and dispersal create fine-scale genetic structure in natural populations — especially of social mammals — with important evolutionary and conservation genetic consequences. Such structure is well-characterized for typical mammalian societies; that is, societies where social group composition is stable, dispersal is male-biased, and males form permanent breeding associations in just one or a few social groups over the course of their lives. However, genetic structure is not well understood for social mammals that differ from this pattern, including elephants. In elephant societies, social groups fission and fuse, and males never form permanent breeding associations with female groups. Here, we combine 33 years of behavioural observations with genetic information for 545 African elephants ( Loxodonta africana ), to investigate how mating and dispersal behaviours structure genetic variation between social groups and across age classes. We found that, like most social mammals, female matrilocality in elephants creates co-ancestry within core social groups and significant genetic differentiation between groups (Φ_ST = 0.058). However, unlike typical social mammals, male elephants do not bias reproduction towards a limited subset of social groups, and instead breed randomly across the population. As a result, reproductively dominant males mediate gene flow between core groups, which creates cohorts of similar-aged paternal relatives across the population. Because poaching tends to eliminate the oldest elephants from populations, illegal hunting and poaching are likely to erode fine-scale genetic structure. We discuss our results and their evolutionary and conservation genetic implications in the context of other social mammals.     

The role of life history in the relationship between population dynamics and environmental variability in two Mediterranean stream fishes

Chub Squalius torgalensis and nase Chondrostoma lusitanicum , in a Mediterranean stream, showed important differences in life-history traits and population dynamics. Both species reached mean maturity at age 2 years. Chub lived up to age 5 years, spawned in March to June, grew at a maximum rate of 0·59 mm mm⁻¹ year⁻¹ and showed a low reproductive allocation, with fecundity and egg size increasing with body size. Nase lived up to age 4 years, spawned in January to April, grew at a maximum rate of 0·46 mm mm⁻¹ year⁻¹ and showed a high reproductive allocation, with egg size independent of body size. Both chub and nase showed moderate fluctuations in population size during 1991–1998, but differed in factors driving density at age. Density of age 1 year juvenile chub decreased following severe summer droughts and proportionate survival prevailed thereafter. Density of age 2 year adult nase decreased following severe spring floods, but neither environmental nor parental stock effects were detected for juveniles and older fishes. The results illustrated the interplay between life history and environmental variability in driving fish population dynamics, with impacts of both summer droughts and spring floods being contingent on species-specific patterns of spawning and reproductive investment.     

Distribution of nucleotide differences between two randomly chosen cistrons in a population of variable size

The distribution of the number of nucleotide differences between two randomly chosen cistrons in a finite population is studied here when the population size changes from generation to generation. When genetic variability is measured by heterozygosity (i.e., the probability that two cistrons are different), by the probability that two cistrons differ at two or more nucleotide sites, or by mean number of site differences between cistrons, it is seen that in a population going through a small bottleneck all of these measures decline rapidly but, as soon as population size becomes large, they start to increase owing to new mutations. The amount of reduction in these measures depends not only on the size of bottleneck but also on the rate of population growth. The implications of this study explaining the observed variations in the rates of amino acid substitutions during the evolutionary process are also discussed.     

A predictive relationship between population and genetic sex ratios in clonal species

Sexual reproduction depends on mate availability that is reflected by local sex ratios. In species where both sexes can clonally expand, the population sex ratio describes the proportion of males, including clonally derived individuals (ramets) in addition to sexually produced individuals (genets). In contrast to population sex ratio that accounts for the overall abundance of the sexes, the genetic sex ratio reflects the relative abundance of genetically unique mates, which is critical in predicting effective population size but is difficult to estimate in the field. While an intuitive positive relationship between population (ramet) sex ratio and genetic (genet) sex ratio is expected, an explicit relationship is unknown. In this study, we determined a mathematical expression in the form of a hyperbola that encompasses a linear to a nonlinear positive relationship between ramet and genet sex ratios. As expected when both sexes clonally have equal number of ramets per genet both sex ratios are identical, and thus ramet sex ratio becomes a linear function of genet sex ratio. Conversely, if sex differences in ramet number occur, this mathematical relationship becomes nonlinear and a discrepancy between the sex ratios amplifies from extreme sex ratios values towards intermediate values. We evaluated our predictions with empirical data that simultaneously quantified ramet and genet sex ratios in populations of several species. We found that the data support the predicted positive nonlinear relationship, indicating sex differences in ramet number across populations. However, some data may also fit the null model, which suggests that sex differences in ramet number were not extensive, or the number of populations was too small to capture the curvature of the nonlinear relationship. Data with lack of fit suggest the presence of factors capable of weakening the positive relationship between the sex ratios. Advantages of this model include predicting genet sex ratio using population sex ratios given known sex differences in ramet number, and detecting sex differences in ramet number among populations.     

Variance effective population size is affected by census size in sub-structured populations

Measurement of allele frequency shifts between temporally spaced samples has long been used for assessment of effective population size (N_e), and this ‘temporal method’ provides estimates of N_e referred to as variance effective size (N_eV). We show that N_eV of a local population that belongs to a sub-structured population (a metapopulation) is determined not only by genetic drift and migration rate (m), but also by the census size (N_c). The realized N_eV of a local population can either increase or decrease with increasing m, depending on the relationship between N_e and N_c in isolation. This is shown by explicit mathematical expressions for the factors affecting N_eV derived for an island model of migration. We verify analytical results using high-resolution computer simulations, and show that the phenomenon is not restricted to the island model migration pattern. The effect of N_c on the realized N_eV of a local subpopulation is most pronounced at high migration rates. We show that N_c only affects local N_eV, whereas N_eV for the metapopulation as a whole, inbreeding (N_eI), and linkage disequilibrium (N_eLD) effective size are all independent of N_c. Our results provide a possible explanation to the large variation of N_e/N_c ratios reported in the literature, where N_e is frequently estimated by N_eV. They are also important for the interpretation of empirical N_e estimates in genetic management where local N_eV is often used as a substitute for inbreeding effective size, and we suggest an increased focus on metapopulation N_eV as a proxy for N_eI.     

The genetic balance between varying population size and selective neutrality

This note extends to an arbitrary offspring distribution the generalized model for random fluctuation of allele frequency, where population size is permitted to fluctuate randomly from generation to generation. Martingale methods analogous to those of Seneta (1974) and Heyde and Seneta (1975) are applied to discuss conditions for Pr(Y(1–Y)>0)>0, where Y is the (almost sure) limiting frequency of one allele. An overlapping generation study of the same phenomenon has recently been made by Heyde (1981).     

The genetic effective and adult census size of an Australian population of tiger prawns (Penaeus esculentus)

This study compares estimates of the census size of the spawning population with genetic estimates of effective current and long-term population size for an abundant and commercially important marine invertebrate, the brown tiger prawn (Penaeus esculentus). Our aim was to focus on the relationship between genetic effective and census size that may provide a source of information for viability analyses of naturally occurring populations. Samples were taken in 2001, 2002 and 2003 from a population on the east coast of Australia and temporal allelic variation was measured at eight polymorphic microsatellite loci. Moments-based and maximum-likelihood estimates of current genetic effective population size ranged from 797 to 1304. The mean long-term genetic effective population size was 9968. Although small for a large population, the effective population size estimates were above the threshold where genetic diversity is lost at neutral alleles through drift or inbreeding. Simulation studies correctly predicted that under these experimental conditions the genetic estimates would have non-infinite upper confidence limits and revealed they might be overestimates of the true size. We also show that estimates of mortality and variance in family size may be derived from data on average fecundity, current genetic effective and census spawning population size, assuming effective population size is equivalent to the number of breeders. This work confirms that it is feasible to obtain accurate estimates of current genetic effective population size for abundant Type III species using existing genetic marker technology.     

Fine-scale genetic population structure of an understory rainforest bird in Costa Rica

We studied five populations of a rainforest understory insectivorous bird (Myrmeciza exsul, chestnut-backed antbird) in a fragmented landscape in northeastern Costa Rica in order to test hypotheses about the influence of forest fragmentation on population genetic structure using 16 microsatellite loci. Bayesian assignment approaches—perhaps the most conservative analyses we performed—consistently grouped the sites into two distinct groups, with all individuals from the smallest and most isolated population clustering separately from the other four sites. Additional analyses revealed (1) overall significant genetic structure; (2) a pattern of population differentiation consistent with a hypothesis of isolation by resistance (landscape connectivity), but not distance; and (3) relatively short dispersal distances indicated by elevated mean pairwise relatedness in several of the sites. Our results are somewhat surprising given the small geographic distances between sites (11–34?km) and the short time (~60?years) since wide-spread deforestation in this landscape. We suspect fine-scale genetic structure may occur in many resident tropical bird species, and in the case of the chestnut-backed antbird it appears that anthropogenic habitat fragmentation has important population genetic implications. It appears that chestnut-backed antbirds may persist in fragmented landscapes in the absence of significant migration among patches, but mechanisms that allow this species to persist when many other similar species do not are not well understood.     

Capwire: a R package for estimating population census size from non‐invasive genetic sampling

Non‐invasive genetic sampling is an increasingly popular approach for investigating the demographics of natural populations. This has also become a useful tool for managers and conservation biologists, especially for those species for which traditional mark–recapture studies are not practical. However, the consequence of collecting DNA indirectly is that an individual may be sampled multiple times per sampling session. This requires alternative statistical approaches to those used in traditional mark–recapture studies. Here we present the R package capwire , an implementation of the population size estimators of Miller et al. (Molecular Ecology 2005; 14 : 1991), which were designed to deal specifically with this type of sampling. The aim of this project is to enable users across platforms to easily manipulate their data and interact with existing R packages. We have also provided functions to simulate data under a variety of scenarios to allow for rigorous testing of the robustness of the method and to facilitate further development of this approach.     

Food relations between coexisting native Hawaiian stream fishes

In this study, an ecomorphological perspective is used to examine the role of feeding morphology in shaping patterns of food resource use and coexistence for Awaous guamensis and Sicyopterus stimpsoni, two native gobies (Gobiidae) which inhabit mountainous streams of the high Hawaiian Islands. Using data from underwater census, gut content analysis, and benthic sampling, I determined that A. guamensis, the generalist, had nearly unchallenged use of invertebrate foods. Overlap in fitness for algal use, however, resulted in a partitioning of benthic algae, with A. guamensis having domain over most green algae (43.0% of diet), whereas S. stimpsoni the algal specialist, fed predominantly on blue-green algae (22.6% of diet) and diatoms (54.2% of diet). Cladophora sp. (Chlorophyta) and pennate diatoms (Chrysophyta) were determined to be the primary algal foods of A. guamensis and S. stimpsoni, respectively, and were utilized in a mutually exclusive manner with other secondary algal foods depending upon availability. Heterogeneity, found in the abundance and composition of algal and invertebrate foods in the benthic landscape both spatially and seasonally, may be regulated by stream flow and periodic disturbance. This changing mosaic of foods is suggested as having provided opportunities for minimizing competitive conflicts and enhancing the potential for stream species to coexist. Competition for preferred foods, created by inter-specific overlap in ecomorphology and spurred by constraints placed on food diversity by the extreme geographic isolation of the Hawaiian Islands, is hypothesized as having played an evolutionary role in shaping resource use patterns which facilitate coexistence.