EVIDENCE THAT AN OUTCROSSING POPULATION IS A DERIVED LINEAGE IN A HERMAPHRODITIC FISH (RIVULUS MARMORATUS)

Rivulus marmoratus is the only known vertebrate with obligate, synchronous hermaphroditic fertilization. Males can be experimentally induced in the laboratory and are rare or absent in most populations, but at the isolated Twin Cays, Belize, locality, males are relatively abundant. At this locality, evidence of outcrossing has been documented in this otherwise automictic cloning species. Phylogenetic analysis of restriction sites and sequence characters revealed that all Florida and Belize western Caribbean populations (including Twin Cays) are phyletically indistinguishable yet divergent from eastern populations in Brazil and the Bahamas. Further, these western lineages shared a common ancestor more recently than all other populations. Therefore, the Twin Cays population is not a remnant ancestral outcrossing population. Outcrossing is suspected to have evolved as a phenotypically plastic character, and its expression in R. marmoratus may be dormant unless triggered by some ecological factor that is not well understood.     

Among- and within-population variation in outcrossing rate of a mixed-mating freshwater snail

Mixed-mating animals self-fertilize a proportion of their offspring. Outcrossing rate may covary with the ecological and historical factors affecting the population. Theory predicts that outcrossing is favored when inbreeding depression is high and when individual heterozygosity is important. Self-fertilization is predicted to be favored when costs of male function, or mate finding are high, for example, when empty patches are colonized by few individuals. In this study, we assessed primary (after hatching) and secondary (after juvenile mortality) outcrossing rates of two mixed-mating snail populations. Our purpose was to assess the variation in mating-system parameters and estimate significance of inbreeding depression for secondary outcrossing rate (the realized outcrossing rate of parents that produce the next generation). Secondary outcrossing rate was higher than the primary outcrossing rate in one of the two populations, suggesting considerable inbreeding depression. In the other study population, secondary outcrossing rates were found to increase when initially low, or decrease when initially high, depending on the family. Moderate outcrossing rates were found to be more stable. Parental inbreeding coefficients were close to zero in both populations. Outcrossing rate was much more variable among families in the population with the lower average outcrossing rate, suggesting that individuals differed considerably in their mating system. Our results add to recent studies suggesting that populations of mixed-mating animals may differ in their mating system parameters and expression of inbreeding depression.     

Assessing the impact of ancient Maya animal use

Tropical forest animals are at high risk worldwide as a result of over-exploitation and forest clearing. Zooarchaeological studies of animal use by the ancient Maya of the southern lowland regions of Guatemala, Honduras, Belize, and Mexico provide long-term historical information on animal populations under conditions of human population growth and climatic change that is valuable to both archaeology and conservation biology. In this paper, zooarchaeological data from 35 chronologically defined faunal sub-samples recovered from 25 ancient Maya archaeological sites are used to assess the effects of ancient hunting on animal populations of the Maya region between the Preclassic and Colonial periods (2000 BC–AD 1697). The variations in species abundance are used as a proxy for describing changes in ancient Maya hunting practices and hunted animal populations, interpreted on the basis of hunting efficiency models from foraging ecology. A significant reduction in the proportion of large mammals, particularly Odocoileus virginianus, in zooarchaeological assemblages between the Late Classic (AD 600–850) and Terminal Classic/Postclassic periods (AD 850–1519) suggest that over-hunting during the Late Classic may have led to a reduction in availability of these animals to the ancient Maya hunters in the later periods. This finding is discussed in relation to important social and environmental variations to evaluate the impact of hunting and other factors such as forest clearance and climate on ancient animal populations in the Maya region.     

Outcrossing rates and male sterility in natural populations of Plantago coronopus

Summary Outcrossing rates were estimated in three populations of the gynodioecious species Plantago coronopus by means of electrophoresis of adult plants and their natural progenies. A multilocus estimation procedure was used. Heterogeneity among the pollen-pool allele frequencies did not exist either in space of in time. Differences between populations in mean outcrossing rates were large (range: 0.34–0.93), probably caused by differences in densities of flowering plants. In addition, there was considerable variability between individuals, which was at least partly caused by the presence of male sterility. Population density may, via its influence on outcrossing rates, be a factor influencing the maintenance of male sterile plants in the population. The level of outcrossing in hermaphrodites was not low enough to explain the maintenance of male steriles. Outcrossing rates in two populations, established via progeny analysis, were much lower than calculated with the fixation index, possibly indicating heterozygote advantage in these natural populations.Grassland Species Research Group Publication no. 134     

Persistence of Mhc heterozygosity in homozygous clonal killifish,Rivulus marmoratus: implications for the origin of hermaphroditism

The mangrove killifish Rivulus marmoratus, a neotropical fish in the order Cyprinodontiformes, is the only known obligatorily selfing, synchronous hermaphroditic vertebrate. To shed light on its population structure and the origin of hermaphroditism, major histocompatibility complex (Mhc) class I genes of the killifish from seven different localities in Florida, Belize, and the Bahamas were cloned and sequenced. Thirteen loci and their alleles were identified and classified into eight groups. The loci apparently arose approximately 20 million years ago (MYA) by gene duplications from a single common progenitor in the ancestors of R. marmoratus and its closest relatives. Distinct loci were found to be restricted to different populations and different individuals in the same population. Up to 44% of the fish were heterozygotes at Mhc loci, as compared to near homozygosity at non-Mhc loci. Large genetic distances between some of the Mhc alleles revealed the presence of ancestral allelic lineages. Computer simulation designed to explain these findings indicated that selfing is incomplete in R. marmoratus populations, that Mhc allelic lineages must have diverged before the onset of selfing, and that the hermaphroditism arose in a population containing multiple ancestral Mhc lineages. A model is proposed in which hermaphroditism arose stage-wise by mutations, each of which spread through the entire population and was fixed independently in the emerging clones.     

Barriers to outcrossing success in the primarily self-fertilizing clam shrimp, Eulimnadia texana (Crustacea, Branchiopoda)

Abstract. The expected proportion of males in androdioecious populations (those comprised of males and hermaphrodites) largely depends on the fertilization opportunities of males. If male mating opportunities are low due to restricted access to hermaphroditic eggs, then populations will be hermaphrodite-biased. Hermaphrodites have two mechanisms available to limit male mating success: (1) pre-mating barriers to outcrossing, in which hermaphrodites choose not to pair with males and (2) post-mating barriers to outcrossing, in which hermaphrodite sperm has greater access to eggs than male sperm. In this study, we measured male mating success in the androdioecious clam shrimp Eulimnadia texana when pre-mating barriers to outcrossing were removed. These branchiopod crustaceans are small (5–8 mm), filter feeders that live in ephemeral pools in the deserts of the southwestern United States. Using genetic markers, we measured male mating success in laboratory experiments in two populations of these shrimp. We correlated mating success with clasping time, clasping during egg transfer, and male thrusting during egg transfer. Males fertilized an average of 24–40% of the hermaphrodites' eggs. Outcrossing success was positively correlated with clasping duration, and was nearly an order of magnitude higher for males thrusting during egg transfer relative to thrusting at other times during pairing. Because these estimates of mating success were similar to previously reported estimates (in which both pre- and post-mating barriers to outcrossing were potentially important), we deduced that pre-mating barriers to outcrossing do not greatly decrease male outcrossing success in E. texana ; the low fertilization (25–50% of available eggs) by males is thus due to post-mating barrier(s) to outcrossing.     

Strong population structure despite evidence of recent migration in a selfing hermaphroditic vertebrate, the mangrove killifish (Kryptolebias marmoratus)

We employ a battery of 33 polymorphic microsatellite loci to describe geographical population structure of the mangrove killifish (Kryptolebias marmoratus), the only vertebrate species known to have a mixed-mating system of selfing and outcrossing. Significant population genetic structure was detected at spatial scales ranging from tens to hundreds of kilometres in Florida, Belize, and the Bahamas. The wealth of genotypic information, coupled with the highly inbred nature of most killifish lineages due to predominant selfing, also permitted treatments of individual fish as units of analysis. Genetic clustering algorithms, neighbour-joining trees, factorial correspondence, and related methods all earmarked particular killifish specimens as products of recent outcross events that could often be provisionally linked to specific migration events. Although mutation is the ultimate source of genetic diversity in K. marmoratus, our data indicate that interlocality dispersal and outcross-mediated genetic recombination (and probably genetic drift also) play key proximate roles in the local 'clonal' dynamics of this species.     

Ecology of spawning humpback chub, Gila cypha, in the Little Colorado River near Grand Canyon, Arizona

The morphologically unique and endangered humpback chub, Gila cypha, is found in canyon-bound reaches of the Colorado River and its tributaries. Now limited to six isolated reproducing populations, this species is believed to have been once distributed over a large portion of the mainstem river. Because the species inhabits remote canyon areas, little is known about its spawning ecology. The largest remaining population occurs in the lower Little Colorado River (LCR) near Grand Canyon, where we conducted a three-year study of spawning ecology during spring (March-June) 1993–1995. We analyzed seasonal patterns of movement, population density, relative condition, spawning scores, and frequency of ripe condition and fin abrasions and compared these data with seasonal discharge and water temperature to determine spawning phenology and ecology. Spawning commenced in late March, peaked in mid-April, and waned in mid-May. A high proportion of males remained ripe over this period, whereas ripe females were relatively abundant only in April. Increased densities of adult fish in March-April and rapid declines in May-June coupled with recaptures of 18.4% of these adults in the Colorado River suggest that a portion of the population migrated from the Colorado River into the LCR to spawn and then returned. Ripe males aggregated in areas of complex habitat structure with high angular variation in bottom profiles (matrix of large boulders, travertine masses combined with chutes, runs and eddies, 0.5–2.0 m deep) and were associated with deposits of clean gravel. Ripe females appeared to move to these male aggregations to spawn. Near-spawning (including gravid) females and non-spawning fish used similar habitats and were segregated but close (< 50 m) to habitats occupied by aggregations of ripe males. Abrasions on anal and lower caudal fins of males and females suggest that spawning involves contact with gravel substrates, where semi-adhesive eggs are deposited and fertilized. The findings of this study should aid recovery efforts for humpback chub by identifying spawning habitat within the historic distributional range where additional spawning stocks could be established.     

Inferring recent outcrossing rates using multilocus individual heterozygosity: application to evolving wheat populations

Using multilocus individual heterozygosity, a method is developed to estimate the outcrossing rates of a population over a few previous generations. Considering that individuals originate either from outcrossing or from n successive selfing generations from an outbred ancestor, a maximum-likelihood (ML) estimator is described that gives estimates of past outcrossing rates in terms of proportions of individuals with different n values. Heterozygosities at several unlinked codominant loci are used to assign n values to each individual. This method also allows a test of whether populations are in inbreeding equilibrium. The estimator's reliability was checked using simulations for different mating histories. We show that this ML estimator can provide estimates of outcrossing rates for the final generation outcrossing rate (t(0)) and a mean of the preceding rates (t(p)) and can detect major temporal variation in the mating system. The method is most efficient for low to intermediate outcrossing levels. Applied to nine populations of wheat, this method gave estimates of t(0) and t(p). These estimates confirmed the absence of outcrossing t(0) = 0 in the two populations subjected to manual selfing. For free-mating wheat populations, it detected lower final generation outcrossing rates t(0) = 0-0.06 than those expected from global heterozygosity t = 0.02-0.09. This estimator appears to be a new and efficient way to describe the multilocus heterozygosity of a population, complementary to Fis and progeny analysis approaches.     

A test of the good-genes-as-heterozygosity hypothesis using red-winged blackbirds

Brown recently proposed that the "good genes" that females pursuewhen choosing mates may be individual heterozygosity becausemore heterozygous mates sire offspring with higher fitness.Further, because heterozygosity might enhance developmentalstability, males with more heterozygosity are recognized bythe reduced fluctuating asymmetry (FA) of their bilaterallypaired traits. We used a point sample of 67 male red-winged blackbirds(Agelaius phoeniceus) to test two predictions of this hypothesis:(1) males with more heterozygosity have higher fitness, and(2) males with more heterozygosity have lower FA. We identified7 polymorphic loci from an initial screening of 16 enzymes;32 individuals were completely homozygous, and 35 individualswere heterozygous at at least 1 locus. Larger and older malesrealized higher mating success in this population, but neither sizenor age was related to heterozygosity. Heterozygous males werenot in better condition than homozygous males, nor were theyless infected by hematozoa, lice, or mites. Among 1-year oldmales, epaulet length did not differ between homozygotes andheterozygotes, but among older males, heterozygotes did havelonger epaulets. Homozygotes and heterozygotes did not differin their mean FA scores for nine individual characters. Althoughthe two groups of males did differ in composite FA, heterozygousmales were less symmetrical. Interestingly, this differencewas attributable to a single allele at the PGM-3 locus. Combinedwith previous results showing that FA was generally unrelatedto male health, viability, parental care, social dominance,or mating success, the present results indicate that Brown's hypothesisdoes not explain mate choice or male quality in this populationof red-winged blackbirds.     

Balance between inbreeding and outcrossing in a nannandrous species,the moss Homalothecium lutescens

Epiphytic dwarf males on the females present a possible solution to the problem of short fertilization distances in mosses. However, leptokurtic spore dispersal makes dwarf males likely to be closely related to their host shoot, with an accompanying risk of inbreeding. The capacity of a female to harbour a high number of different dwarf males suggests that there may be mechanisms in place that counteract inbreeding, such as polyandry and post-fertilization selection. We have genotyped sporophytes, female host shoots and dwarf males in four populations of the moss Homalothecium lutescens. We found no evidence of selective sporophyte abortion based on level of heterozygosity. The occurrence of entirely homozygous sporophytes together with significantly positive inbreeding coefficients in three of the populations (mean F_IS between 0.48 and 0.64) suggest frequent mother–son mating events. However, 23% of all sampled sporophytes had a higher level of heterozygosity compared with the mean expected heterozygosity at the population level. Polyandry was frequent, on average 59% of the sporophytes on a female shoot were sired by distinct fathers. In conclusion, sporadic fertilizations by dwarf males originating from nonhost female shoots appear to counteract strong inbreeding.     

Increased inbreeding but not homozygosity in small populations of Sabatia angularis (Gentianaceae)

Understanding how the mating system varies with population size in plant populations is critical for understanding their genetic and demographic fates. We examined how the mating system, characterized by outcrossing rate, biparental inbreeding rate, and inbreeding coefficient, and genetic diversity varied with population size in natural populations of the biennial Sabatia angularis. We found a significant, positive relationship between outcrossing and population size. Selfing was as high as 40% in one small population but was only 7% in the largest population. Despite this pattern, observed heterozygosity did not vary with population size, and we suggest that selection against inbred individuals maintains observed heterozygosity in small populations. Consistent with this hypothesis, we found a trend of lower inbreeding coefficients in the maternal than progeny generation in all of the populations, and half of the populations exhibited significant excesses of adult heterozygosity. Moreover, genetic diversity was not related to population size and was similar across all populations examined. Our results suggest that the consequences of increased selfing for population fitness in S. angularis, a species that experiences significant inbreeding depression, will depend on the relative magnitude and consistency of inbreeding depression and the demographic cost of selection for outcrossed progeny in small populations.     

Genetic Evidence for Multiple Sources of the Non-Native Fish Cichlasoma urophthalmus (Günther; Mayan Cichlids) in Southern Florida

The number and diversity of source populations may influence the genetic diversity of newly introduced populations and affect the likelihood of their establishment and spread. We used the cytochrome b mitochondrial gene and nuclear microsatellite loci to identify the sources of a successful invader in southern Florida, USA, Cichlasoma urophthalmus (Mayan cichlid). Our cytochrome b data supported an introduction from Guatemala, while our microsatellite data suggested movement of Mayan Cichlids from the upper Yucatán Peninsula to Guatemala and introductions from Guatemala and Belize to Florida. The mismatch between mitochondrial and nuclear genomes suggests admixture of a female lineage from Guatemala, where all individuals were fixed for the mitochondrial haplotype found in the introduced population, and a more diverse but also relatively small number of individuals from Belize. The Florida cytochrome b haplotype appears to be absent from Belize (0 out of 136 fish screened from Belize had this haplotype). Genetic structure within the Florida population was minimal, indicating a panmictic population, while Mexican and Central American samples displayed more genetic subdivision. Individuals from the Upper Yucatán Peninsula and the Petén region of Guatemala were more genetically similar to each other than to fish from nearby sites and movement of Mayan Cichlids between these regions occurred thousands of generations ago, suggestive of pre-Columbian human transportation of Mayan Cichlids through this region. Mayan Cichlids present a rare example of cytonuclear disequilibrium and reduced genetic diversity in the introduced population that persists more than 30 years (at least 7–8 generations) after introduction. We suggest that hybridization occurred in ornamental fish farms in Florida and may contribute their establishment in the novel habitat. Hybridization prior to release may contribute to other successful invasions.     

Discrimination of French grunts (Haemulon flavolineatum, Desmarest, 1823) from mangrove and coral reef habitats using otolith microchemistry

Using French Grunts (Haemulon flavolineatum) held captive within mangrove and reef sites, we determined (a) whether otolith microchemical differences existed between mangroves and reefs separated at a biologically relevant spatial scale (0.25-7.1 km), (b) whether patterns in elemental concentrations were consistent across years, and (c) whether it was possible to identify whether a given fish occupied a mangrove as a juvenile.Three sites were established at Great Exuma, Bahamas (two reefs and one mangrove, May 2001) and at Turneffe Atoll, Belize (one reef and two mangroves, August 2001 and 2002). Using concentrations of Sr and Ba, discriminant function analysis (DFA) indicated unique spatial microchemical signatures of fish from each of the three Bahamas sites allowing an average correct classification of 77%. Using concentrations of Sr, Ba, Sn and Pb (2001), and in addition Li, Mg, Cu and Rb (2002), DFA of the three Belize sites indicated an average correct classification of 68% and 85% in 2001 and 2002, respectively. To assess temporal variability in otolith microchemistry, we compared microchemical signatures of Belize fish from 2001 to those from 2002. On average, 42% of fish from 2002 were correctly classified to their captive sites using chemical information from 2001, thus suggesting considerable temporal variability in otolith microchemistry. Finally, to identify whether a given fish occupied a mangrove during its juvenile stage, we ablated the juvenile portion of the otolith taken from reefs in Belize 2002. Results of this analysis indicated that 36% of 39 individuals taken from the reef had a signature more representative of one of the mangrove sites. Although otolith microchemistry varied temporally and our analysis was restricted to the grouping of individuals to only one of three sites, mangroves appeared to contribute to reef populations.     

BREEDING SYSTEM EVOLUTION IN LEAVENWORTHIA: BREEDING SYSTEM VARIATION AND REPRODUCTIVE SUCCESS IN NATURAL POPULATIONS OF LEAVENWORTHIA CRASSA (CRUCIFERAE)

Populations of Leavenworthia crassa (Cruciferae) studied for 3 years exhibited among- and within-population genetic variation for a suite of floral and reproductive traits (flower width, petal length, anther position, ability to set seeds in the absence of pollinators, time to first flowering) associated with breeding system. We used electrophoretic markers to show that a population with small, monomorphically colored flowers with introrse anthers had a significantly lower outcrossing rate (t = 0.03) than did a population with larger, polymorphically colored flowers with extrorse anthers (t = 0.33). In the more-outcrossing population the correlation between higher maternal plant outcrossing rate and the suite of six traits approached significance (P < 0.067), with greater petal size, greater flower width, and reduced ability to set seeds in the absence of pollinators contributing significantly. Plants in selfing populations had a generally higher reproductive success, with a higher number of flowers per plant, a smaller proportion of unfertilized ovules, a smaller proportion of fertilized ovules aborted, a higher rate of fruit set, and overall a larger number of seeds matured than did plants from the more outcrossing populations. Pollen limitation did not appear to account for lower reproductive success in outcrossing populations. Resource limitation did not differ substantially between populations. However, within-ovary patterns of fertilization, abortion, and seed weight were significantly less random in outcrossing populations than in selfing populations, suggesting that differential gamete and embryo success may be responsible for lower reproductive success in outcrossing populations.     

Maintenance of androdioecy in the freshwater shrimp, Eulimnadia texana: estimates of inbreeding depression in two populations

Androdioecy is an uncommon form of reproduction in which males coexist with hermaphrodites. Androdioecy is thought to be difficult to evolve in species that regularly inbreed. The freshwater shrimp Eulimnadia texana has recently been described as both androdioecious and highly selfing and is thus anomalous. Inbreeding depression is one factor that may maintain males in these populations. Here we examine the extent of "late" inbreeding depression (after sexual maturity) in these clam shrimp using two tests: (1) comparing the fitness of shrimp varying in their levels of individual heterozygosity from two natural populations that differ in overall genetic diversity; and (2) specifically outcrossing and selfing shrimp from these same populations and comparing fitness of the resulting offspring. The effects of inbreeding differed within each population. In the more genetically diverse population, fecundity, size, and mortality were significantly reduced in inbred shrimp. In the less genetically diverse population, none of the fitness measures was significantly lowered in selfed shrimp. Combining estimates of early inbreeding depression from a previous study with current estimates of late inbreeding depression suggests that inbreeding depression is substantial (delta = 0.68) in the more diverse population and somewhat lower (delta = 0.50) in the less diverse population. However, given that males have higher mortality rates than hermaphrodites, neither estimate of inbreeding depression is large enough to account for the maintenance of males in either population by inbreeding depression alone. Thus, the stability of androdioecy in this system is likely only if hermaphrodites are unable to self-fertilize many of their own eggs when not mated to a male or if male mating success is generally high (or at least high when males are rare). Patterns of fitness responses in the two populations were consistent with the hypothesis that inbreeding depression is caused by partially recessive deleterious alleles, although a formal test of this hypothesis still needs to be conducted.     

Sex differences in host defence interfere with parasite‐mediated selection for outcrossing during host–parasite coevolution

The Red Queen hypothesis proposes that coevolving parasites select for outcrossing in the host. Outcrossing relies on males, which often show lower immune investment due to, for example, sexual selection. Here, we demonstrate that such sex differences in immunity interfere with parasite‐mediated selection for outcrossing. Two independent coevolution experiments with Caenorhabditis elegans and its microparasite Bacillus thuringiensis produced decreased yet stable frequencies of outcrossing male hosts. A subsequent systematic analysis verified that male C. elegans suffered from a direct selective disadvantage under parasite pressure (i.e. lower resistance, decreased sexual activity, increased escape behaviour), which can reduce outcrossing and thus male frequencies. At the same time, males offered an indirect selective benefit, because male‐mediated outcrossing increased offspring resistance, thus favouring male persistence in the evolving populations. As sex differences in immunity are widespread, such interference of opposing selective constraints is likely of central importance during host adaptation to a coevolving parasite.     

Evolution of outcrossing in experimental populations of Caenorhabditis elegans

Caenorhabditis elegans can reproduce exclusively by self-fertilization. Yet, males can be maintained in laboratory populations, a phenomenon that continues to puzzle biologists. In this study we evaluated the role of males in facilitating adaptation to novel environments. For this, we contrasted the evolution of a fitness component exclusive to outcrossing in experimental populations of different mating systems. We introgressed a modifier of outcrossing into a hybrid population derived from several wild isolates to transform the wild-type androdioecious mating system into a dioecious mating system. By genotyping 375 single-nucleotide polymorphisms we show that the two populations had similar standing genetic diversity available for adaptation, despite the occurrence of selection during their derivation. We then performed replicated experimental evolution under the two mating systems from starting conditions of either high or low levels of diversity, under defined environmental conditions of discrete non-overlapping generations, constant density at high population sizes (N = 10⁴), no obvious spatial structure and abundant food resources. During 100 generations measurements of sex ratios and male competitive performance showed: 1) adaptation to the novel environment; 2) directional selection on male frequency under androdioecy; 3) optimal outcrossing rates of 0.5 under androdioecy; 4) the existence of initial inbreeding depression; and finally 5) that the strength of directional selection on male competitive performance does not depend on male frequencies. Taken together, these results suggest that androdioecious males are maintained at intermediate frequencies because outcrossing is adaptive.     

Reproductive biology and genetic structure in Lloydia serotina

Concern regarding the conservation status of small, isolated populations of the arctic-alpine plant species Lloydia serotina prompted research to establish the status and performance of this species in Wales, in comparison with large populations in its more typical alpine habitat. Relationships between reproductive strategies and genetic variation were investigated in a number of populations, representing a wide habitat, geographic and population size range. In all populations, vegetative reproduction predominates over sexual reproduction, but seed produced is viable and germinates readily under controlled conditions. Smaller, peripheral populations produced fewer flowers and seeds than the larger ones, but all populations studied supported significant percentages (>30%) of male plants, due to either the occurrence of androdioecy in this species or to a resource limited breeding system. Analysis of allozyme variation in sixteen populations from North America, the European Alps and Wales showed lower levels of genetic variation in smaller populations which averaged 1.1–1.2 alleles per locus and 10–20% of loci polymorphic, whereas larger populations averaged 1.4 alleles per locus and 30–40% polymorphic loci. This applied especially to the most northerly and southerly populations in North America, suggesting the occurrence of genetic drift in these small, peripheral populations. F-statistics suggest relatively high levels of differentiation among smaller populations, even among those closely related geographically, but genetic variation has been retained in all but one population, possibly due to infrequent sexual reproduction by long lived clones. RAPD analysis of four small populations in Wales provided further evidence of clonal growth and possible inbreeding dominating a mixed mating reproductive system with consequent genetic structuring in these populations.     

Effects of population outcrossing on rotifer fitness

Background  

Outcrossing between populations can exert either positive or negative effects on offspring fitness. Cyclically parthenogenetic rotifers, like other continental zooplankters, show high genetic differentiation despite their high potential for passive dispersal. Within this context, the effects of outcrossing may be relevant in modulating gene flow between populations through selection for or against interpopulation hybrids. Nevertheless, these effects remain practically unexplored in rotifers. Here, the consequences of outcrossing on the rotifer Brachionus plicatilis were investigated. Cross-mating experiments were performed between a reference population and three alternative populations that differed in their genetic distance with regard to the former. Two offspring generations were obtained: F1 and BC ('backcross'). Fitness of the outcrossed offspring was compared with fitness of the offspring of the reference population for both generations and for three different between-population combinations. Four fitness components were measured throughout the rotifer life cycle: the diapausing egg-hatching proportion, clone viability (for the clones originating from diapausing eggs), initial net growth rate R for each viable clone, and the proportion of male-producing clones. Additionally, both the parental fertilisation proportion and a compound fitness measure, integrating the complete life cycle, were estimated.