Polymorphism of a genital organ under sexual selection in Monacha kuznetsovi from the Caucasus (Gastropoda: Hygromiidae)

During mating the dart apparatus of the land snail group Helicoidea transfers an allohormone that increases paternity success. Thus, this organ is under sexual selection. Despite this selective advantage, the dart sac has independently been lost in many lineages in the Helicoidea, presumably because natural selection counteracts sexual selection. However, the selective pressures that result in the loss of the organ have not been examined experimentally because most species show no variation in the presence or absence of the dart sac. Using nuclear multilocus data and mitochondrial sequences, we show that a so far misidentified morph of the genus Monacha without appendicula, a homologue of the dart sac, is conspecific with M. kuznetsovi with appendicula. This is only the second case of polymorphy with regard to the presence or absence of the dart sac or its homologue in the Helicoidea. Monacha kuznetsovi might therefore be a suitable model organism to study how the interplay between sexual selection and natural selection may affect the evolution of the genital organs.     

EVIDENCE FOR PREDATION AS A FACTOR IN DETERMINING SHELL COLOR FREQUENCIES IN A MANGROVE SNAIL LITTORINA SP. (PROSOBRANCHIA: LITTORINIDAE)

It was hypothesized that in Littorina populations living on Avicennia marina in Moreton Bay, Queensland, yellow shelled individuals are at a selective advantage over other shell colors and that this advantage is due to differential selection by predators. Yellow shelled individuals were more likely to be recaptured than others, indicating a higher survival rate of yellows. When predation was restricted on ten mangrove trees, the apparent advantage of yellow shells was removed. After 18 months, the relative frequency of yellow shelled individuals was significantly lower on experimental trees than on control trees. A combination of selection for crypsis and for the less common morph is suggested as the mechanism maintaining the high levels of variation in this species.     

Genetic markers in the study of Anisakis typica (Diesing, 1860): larval identification and genetic relationships with other species of Anisakis Dujardin, 1845 (Nematoda: Anisakidae)

Genetic variation at 21 gene-enzyme systems was studied in a sample of an adult population of Anisakis typica (Diesing, 1860) recovered in the dolphin Sotalia fluviatilis from the Atlantic coast of Brazil. The characteristic alleles, detected in this population, made it possible to identify as A. typica, Anisakis larvae with a Type I morphology (sensu Berland, 1961) from various fishes: Thunnus thynnus and Auxis thazard from Brazil waters, Trachurus picturatus and Scomber japonicus from Madeiran waters, Scomberomorus commerson, Euthynnus affinis, Sarda orientalis and Coryphaena hippurus from the Somali coast of the Indian Ocean, and Merluccius merluccius from the Eastern Mediterranean. Characteristic allozymes are given for the identification, at any life-stage and in both sexes, of A. typica and the other Anisakis species so far studied genetically. The distribution of A. typica in warmer temperate and tropical waters is confirmed; the definitive hosts so far identified for this species belong to delphinids, phocoenids and pontoporids. The present findings represent the first established records of intermediate/paratenic hosts of A. typica and extend its range to Somali waters of the Indian Ocean and to the Eastern Mediterranean Sea. A remarkable genetic homogeneity was observed in larval and adult samples of A. typica despite their different geographical origin; interpopulation genetic distances were low, ranging from D _Nei=0.004 (Eastern Mediterranean versus Somali) to D _Nei=0.010 (Brazilian versus Somali). Accordingly, indirect estimates of gene flow gave a rather high average value of Nm = 6.00. Genetic divergence of A. typica was, on average, D _Nei=1.12 from the members of the A. simplex complex (A. simplex s.s, A. pegreffii, A. simplex C) and D _Nei=1.41 from A. ziphidarum, which all share Type I larvae; higher values were found from both A. physeteris (D _Nei=2.77)     

Pollen competition between morphs in a pollen‐color dimorphic herb and the loss of phenotypic polymorphism within populations

Flower color polymorphism is relatively uncommon in natural flowering plants, suggesting that maintenance of different color morphs within populations is difficult. To address the selective mechanisms shaping pollen‐color dimorphism, pollinator preferences and reproductive performance were studied over three years in Epimedium pubescens in which some populations had plants with either green or yellow pollen (and anthers). Visitation rate and pollen removal and receipt by the bee pollinator (Andrena emeishanica) did not differ between the two color morphs. Compared to the green morph, siring success of the yellow morph's pollen was lower, but that of mixtures of pollen from green and yellow morphs was lowest. This difference, corresponding to in vivo and ex vivo experiments on pollen performance, indicated that pollen germination, rather than tube growth, of the green morph was higher than that of the yellow morph and was seriously constrained in both morphs if a pollen competitor was present. A rare green morph may invade a yellow‐morph population, but the coexistence of pollen color variants is complicated by the reduced siring success of mixed pollinations. Potential pollen competition between morphs may have discouraged the maintenance of multiple phenotypes within populations, a cryptic mechanism of competitive exclusion.     

Sexual Morph of <i>Furcasterigmium furcatum</i> (Plectosphaerellaceae) from <i>Magnolia liliifera</i> Collected in Northern Thailand

We isolated an interesting fungus from dead leaves of Magnolia liliifera collected from Chiang Mai, Thailand. The novel strain is related to Plectosphaerellaceae based on the morphology of its asexual morph and the analysis of sequence data. Phylogenetic analyses using a combined gene analysis of LSU and ITS sequence data showed that this strain is clustered in the same clade with Furcasterigmium furcatum with high statistical support. The new strains produced the asexual morph in culture which is morphologically similar to F. furcatum. Thus, we identified this strain as the sexual morph of F. furcatum. This is the first record of sexual morph for the monotypic genus Furcasterigmium and the first record of this genus on Magnolia.     

Morph‐dependent fitness and directional change of morph frequencies over time in a Dutch population of Common buzzards Buteo buteo

How genetic polymorphisms are maintained in a population is a key question in evolutionary ecology. Previous work on a plumage colour polymorphism in the common buzzard Buteo buteo suggested heterozygote advantage as the mechanism maintaining the co‐existence of three morphs (light, intermediate and dark). We took advantage of 20 years of life‐history data collected in a Dutch population to replicate earlier studies on the relationship between colour morph and fitness in this species. We examined differences between morphs in adult apparent survival, breeding success, annual number of fledglings produced and cumulative reproductive success. We found that cumulative reproductive success differed among morphs, with the intermediate morph having highest fitness. We also found assortative mating for colour morph, whereby assortative pairs were more likely to produce offspring and had longer‐lasting pair bonds than disassortative pairs. Over the 20‐year study period, the proportion of individuals with an intermediate morph increased. This apparent evolutionary change did not just arise from selection on individual phenotypes, but also from fitness benefits of assortative mating. The increased frequency of intermediates might also be due to immigration or drift. We hypothesize that genetic variation is maintained through spatial variation in selection pressures. Further studies should investigate morph‐dependent dispersal behaviour and habitat choice.     

Survival rates in a natural population of the damselfly Ceriagrion tenellum: effects of sex and female phenotype

1. Ceriagrion tenellum females show genetic colour polymorphism. Androchrome (erythrogastrum) females are brightly (male‐like) coloured while gynochrome females (typica and melanogastrum) show cryptic colouration. 2. Several hypotheses have been proposed to explain the existence of more than one female morph in damselfly populations. The reproductive isolation and intraspecific mimicry hypotheses predict greater survival of gynochrome females, while the density dependent hypothesis predicts no differential survival between morphs. 3. Mature males had greater recapture probability than females while the survival probability was similar for both sexes. Survival and recapture rates were similar for androchrome and gynochrome females. 4. Gynochrome females showed greater mortality or migration rate than androchrome females during the pre‐reproductive period. This result is not predicted by the above hypotheses or by the null hypothesis that colour polymorphism is only maintained by random factors: founder effects, genetic drift, and migration.     

Ecological mechanisms for coexistence of colour polymorphism in a coral-reef fish: an experimental evaluation

The evolution of different colour morphs and how they are maintained in animal populations is poorly understood. We investigated the mechanisms maintaining yellow and brown morphs of a coral-reef fish, Pseudochromis fuscus, at Lizard Island, on the Great Barrier Reef. Histological examination of the gonads revealed that colour morphs were not sex-limited, therefore sexual selection does not appear to promote dichromatism in this species. The field distributions of the two colour morphs were spatially segregated, limiting the opportunity for negative frequency-dependent selection to operate. Our results support another ecological mechanism of coexistence. The yellow morph occurred in deeper areas, usually close to the reef edge, where there was a proportionally high cover of live branching corals. In contrast, the brown morph occurred in shallower areas, more distant from the reef edge, that were proportionally low in live branching corals. Within these habitats, each colour morph of P. fuscus displayed a close association with similar coloured damselfishes from the genus Pomacentrus. The yellow morph was associated with predominantly yellow damselfishes (P. moluccensis and P. amboinensis) and the brown morph with darker coloured species (P. adelus and P. chrysurus). Multiple-choice experiments in the laboratory revealed that: (1) each colour morph of P. fuscus preferentially selected habitat patches occupied by damselfishes with the same colouration; and (2) differences in microhabitat use between the two colour morphs of P. fuscus were due to the presence of different coloured damselfishes in these microhabitats. P. fuscus is a predator of newly recruited damselfishes and the striking resemblance between each morph of P. fuscus and the damselfish with which it was associated, suggests that aggressive mimicry may promote coexistence of P. fuscus colour morphs.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Differential winter mortality and seasonal selection in the polymorphic ladybird Adalia bipunctata (L) in the Netherlands

Seasonal selection acting on the melanic polymorphism in the two-spot ladybird Adalia bipunctata was investigated in The Netherlands. An increase in melanic frequency over the spring-summer reproductive period was quantified. The selective advantage gained by melanics averaged 9%, but significant heterogeneity occurred between populations. Adult hibernation behaviour is described. The beetles when outdoors show a highly clumped distribution both between and within trees. The distribution of the morph classes between aggregations is random. Survivorship in a hibernating cohort (initial n= 1898) on a grid of 70 lime trees near Utrecht was monitored by making three counts over the winter of 1981–1982. Intense selection favouring each melanic morph occurred during December and January. The relative fitness of non-melanics was 0.55 (melanics =1). The discovery of dead beetles in late January (about 5% of total losses) and the absence of spatially density-dependent mortality were consistent with a climatic stress rather than selective predation. The period of selection was associated with very cold temperatures averaging up to 4°C below normal and an overall mortality of nearly 75%. There was no change in morph frequency, near normal temperatures and a lower mortality from February to early April. Examination of groups of nearby trees in late January strongly suggested that similar differential mortality had occurred except on some willows. This difference was probably due to the more protected hibernation sites available on these trees. Samples of hibernating cohorts at three other sites showed no evidence of differential mortality. Laboratory experiments with hibernating beetles found no difference in survivorship or rate of weight loss between starved non-melanics and melanics in temperature regimes with and without periods of adult activity. It is concluded that the intense winter selection on the study limes is probably exceptional. Examination of changes in morph frequency through the annual cycle suggests that at some sites the selection favouring melanics during reproduction is counterbalanced by selection against melanics in late summer or early autumn. The results are discussed in relation to mathematical models of cyclical selection and to other field studies including that of Timoféeff-Ressovsky (1940), who found large decreases in melanic frequency during hibernation in Berlin.     

Frequency-dependent morph differences in kleptoparasitic chase rate in the polymorphic arctic skua Stercorarius parasiticus

The relationship between colouration and kleptoparasitic efficiency (success rate and chase rate) was studied in the polymorphic arctic skua, Stercorarius parasiticus, at a Mediterranean migration site. A shift in the ratio of morphs through the spring, presumably caused by differential migration between populations that differ in morph frequency (the higher the latitude, the more frequent is the pale morph), allowed the comparison of the relative performances of pale and dark birds under different scenarios of morph frequency. Chase rate showed a clear frequency-dependent effect in chases by solitary skuas, birds of the rare morph always conducting more chases than expected. This trend was explained by chases of yellow-legged gulls, Larus michahellis, a relatively large and aggressive species that was difficult to chase (i.e. chases resulted in low success rate) but provided potentially higher reward (i.e. more food) than any other host. These differences could result from the perceived cost-benefit balance of chases, as skuas from the rare morph could find an advantage in chasing this host species by being less rapidly recognised. This is in agreement with the apostatic selection hypothesis, which says that the rare morph in a predator or parasite population is favoured through its unfamiliarity to potential prey or hosts (the surprise effect). Differences in success rate could have provided further support to this interpretation, but sample size was not sufficient for a proper comparison at the host-species level.     

Evolution towards dioecy in heterostylous populations

The effective gender of a plant is defined as the proportion of the plant's genes that are transmitted through pollen (its maleness) or through ovules (its femaleness). Formulae are derived that enable the average effective gender of the morphs of a heterostylous population and the gender of individual plants to be estimated. Estimates require a knowledge of the morph ratio and the seed set that results in each morph from self-fertilization and from legitimate and illegitimate cross-fertilizations. If no illegitimate fertilizations occur, the average gender of the morphs can be estimated from their seed production alone; in this situation the average femaleness of one morph is exactly equal to the average maleness of the other morph. The average gender of long- and short-styled morphs is calculated for populations of fiveCordia species (Boraginaceae) from published data ofOpler & al. (1975). In two species, both morphs transmit their genes equally through pollen and ovules. In the other species, the long-styled morph acts predominantly (or in one species exclusively) as an ovule parent and the short-styled morph succeeds predominantly (or exclusively) as a pollen parent.The features of the evolutionary pathway from heterostyly to dioecy and the selective forces that may be responsible are outlined.     

Mate search and aggregation behaviour in the Galician hybrid zone of Littorina saxatilis

In Galician rocky shores two ecotypes of the snail L. saxatilis can be found in sympatry. A ridged and banded ecotype (RB-morph) and a smooth and unbanded ecotype (SU-morph) overlap in midshore with the production of some hybrids. The distinct morphs mate assortatively and there is evidence of a partial reproductive barrier between them. This sexual isolation is caused by a nonrandom microdistribution and mate choice behaviour. Mucus trail-following, movement rate and aggregation behaviour were studied to determine their roles in the mating behaviour and sexual isolation of this species. Morph-specific mucus trail-following could not, in our experiments, explain either of these two processes. The reasons for the aggregation of morphs were investigated by Monte Carlo simulations of data from natural populations, which showed that size aggregation (refuge sizes fit different sized morphs differently) could explain only about 36% of the morph aggregation in adult snails. In the laboratory, morph aggregation was still present, and simulations suggested that size aggregation was the possible explanation. Thus, morph aggregation in Galician L. saxatilis has to be explained also by other causes in addition to size aggregation. These may be a combination of contrasting preferences for barnacle and mussel patches in the two morphs, and possibly longer copulation and pair formation time with similar sized snails of the same morph. Thus aggregation behaviour, but not trail-following, contributes to incipient reproductive isolation and perhaps sympatric speciation in Galician L. saxatilis populations.     

Reproductive costs of Chaoborus-induced polymorphism in Daphnia pulex

Although the Chaoborus-induced spined morph of Daphnia pulex survives attacks by Chaoborus over twice as frequently as the typical morph, the spined morph is never found in the absence of Chaoborus. This implies that a disadvantage is associated with the spined morph in the absence of Chaoborus predation. The present study tested the hypothesis that the typical morph has a higher intrinsic rate of increase than the spined morph, by measuring several life history characteristics in controlled laboratory experiments at constant temperature and unlimited food.The results suggest that the spined morph of D. pulex takes longer to reach maturity, is smaller at maturity, but has similar egg number and egg size as the typical morph. These results are consistent with the hypothesis that the Chaoborus-induced spined morph is reproductively inferior to the typical morph.     

VARIATION IN THE MATING SYSTEM OF EICHHORNIA PANICULATA (SPRENG.) SOLMS. (PONTEDERIACEAE)

A multilocus procedure was used to estimate outcrossing rates from allozyme data in nine populations of Eichhornia paniculata from NE Brazil and Jamaica. The populations were chosen to represent stages in a proposed model of the evolutionary breakdown of tristyly to semi-homostyly; they differed in morph structure (trimorphic, dimorphic, or monomorphic) and floral traits likely to influence the mating system. The interpopulation range in outcrossing rate, t, was 0.96–0.29. Two additional populations from Jamaica, composed exclusively of self-pollinating, semi-homostylous, mid-styled plants, were invariant at 21 isozyme loci, precluding estimates of outcrossing frequency. Trimorphic populations from Brazil had uniformly high outcrossing rates of 0.96–0.88. Values for the floral morphs within populations were not significantly different. A controlled pollination experiment, comparing the competitive ability of self and cross pollen using the Got-3 marker locus, provided evidence that the maintenance of high outcrossing rates in trimorphic populations results from the prepotency of cross pollen and/or the selective abortion of selfed zygotes. Morph-dependent variation in t was detected within a dimorphic population with the L morph outcrossing with a frequency of 0.76 in comparison with 0.36 in the M morph. The difference in the mating system of floral morphs results from modifications in position of short-level stamens in flowers of the M morph resulting in automatic self-pollination. The occurrence of E. paniculata populations composed exclusively of self-pollinating, mid-styled variants is thought to be associated with the spread of genes modifying stamen position. The high level of self-fertilization demonstrated in the M morph would allow automatic selection of these genes, augmented by fertility assurance in the absence of specialized pollinators.     

Resistance of an invasive gastropod to an indigenous trematode parasite in Lake Malawi

Successful establishment and spread of biological invaders may be promoted by the absence of population-regulating enemies such as pathogens, parasites or predators. This may come about when introduced taxa are missing enemies from their native habitats, or through immunity to enemies within invaded habitats. Here we provide field evidence that trematode parasites are absent in a highly invasive morph of the gastropod Melanoides tuberculata in Lake Malawi, and that the invasive morph is resistant to indigenous trematodes that castrate and induce gigantism in native M. tuberculata. Since helminth infections can strongly influence host population abundances in other host-parasite systems, this enemy release may have provided an advantage to the invasive morph in terms of reproductive capacity and survivorship.     

A bird’s eye view of the peppered moth

Industrial melanism in Biston betularia is one of the best known examples of the role of natural selection in evolution and has received considerable scrutiny for many years. The rise in frequency of the dark form of the moth (carbonaria) and a decrease in the pale form (typica) was the result of differential predation by birds, the melanic form being more cyptic than typica in industrial areas where the tree bark was darkened by air pollution. One important aspect of early work evaluating the relative crypsis of the forms of B. betularia on tree trunks with different lichen flora was the reliance on human observers. Humans, however, do not have the same visual capabilities as birds. Birds have well‐developed ultraviolet (UV) vision, an important component of their colour processing system that affects many aspects of behaviour, including prey detection. We examined the UV characteristics of the two forms of B. betularia and a number of foliose and crustose lichens. In human visible light the speckled form typica appeared cyptic when seen against a background of foliose lichen, whereas the dark form carbonaria was conspicuous. Under UV light the situation was reversed. The foliose lichens absorbed UV and appeared dark as did carbonaria. Typica, however, reflected UV and was conspicuous. Against crustose lichens, typica was less visible than carbonaria in both visible and UV light. These findings are considered in relation to the distribution and recolonization of trees by lichens and the resting behaviour of B. betularia.     

Thermal biology of the common garter snake Thamnophis sirtalis (L.)

Summary The thermoregulatory significance of a striped-melanic colour polymorphism in the common garter snake, Thamnophis sirtalis, was assessed through a combination of labortory experimentation and field study. In experiments with living snakes the melanic morph maintained a higher body temperature than the striped morph, when exposed to natural insolation. Experiments with excised skin showed that this thermal advantage is attributable to some integumental difference between the two morphs. Body temperatures of snakes in the field revealed that, during the colder part of the active season, melanics were able to stay warmer than striped snakes by an amount (1.24 C°) approximating the difference observed in the laboratory. Some evidence and argument is presented to suggest that melanism also may confer protection against overheating in warm periods.     

HISTORICAL FACTORS AND ANISOPLETHIC POPULATION STRUCTURE IN TRISTYLOUS PONTEDERIA CORDATA: A REASSESSMENT

Theoretical models of floral-morph frequencies in tristylous species predict a single equilibrium with all three morphs represented in equal proportions (isoplethy). North American populations of Pontederia cordata exhibit considerable heterogeneity of morph frequencies between populations, with the short-styled morph often in excess of isoplethic expectations and the long-styled morph commonly underrepresented. In a previous study, it was proposed that anisoplethic population structure in P. cordata is the result of differential male fertility, owing to genetic differences in pollen production among the morphs. In this study, the influence of historical factors on morph frequencies prior to equilibrium was investigated using a deterministic computer model. Nonequilibrium frequencies are strongly influenced by the genotypes of founding individuals, and, because tristyly is under the control of two diallelic loci, phenotypic equilibrium is approached asymptotically. The model indicates that in nonequilibrium populations the short-styled morph will be in excess and the long-styled morph will be underrepresented. This suggests that historical factors play an important role in determining population structure in P. cordata. Several features of the population ecology of the species lend support to this interpretation. Historical factors should be taken into account when interpreting data from population surveys of morph frequencies in tristylous species and of other genetic polymorphisms not under single-locus control.     

EXPERIMENTAL STUDIES ON THE FUNCTIONAL SIGNIFICANCE OF HETEROSTYLY

Heterostyly has been viewed as both an antiselfing device and a mechanism that increases the proficiency of pollen transfer between plants. We used experimental manipulation of the morph structure of garden populations of self-compatible, tristylous Eichhornia paniculata to investigate the function of floral polymorphism. Outcrossing rates (t), levels of intermorph mating (d), and morph-specific male and female reproductive success were compared in replicate trimorphic and monomorphic populations. In trimorphic populations, t and d averaged 0.81 (2 SE = 0.03) and 0.77 (2 SE = 0.03) respectively, with no difference in either parameter among morphs. Ninety-five percent of outcrossed seeds were therefore the result of intermorph fertilizations. Male reproductive success of the long-styled morph was low, especially in comparison with plants of the short-styled morph. Outcrossing rates for each morph were higher in trimorphic than monomorphic populations where t averaged 0.71 (2 SE = 0.01), 0.30 (2 SE = 0.04) and 0.43 (2 SE = 0.1) for the long-, mid-, and short-styled morphs, respectively. Seed set was lower in monomorphic populations, particularly those composed of the L morph, reflecting reduced pollen deposition. Floral polymorphism therefore increased both outcrossing rate and fecundity but the magnitude of the differences varied among morphs. If the ancestral condition in heterostylous groups resembled the L morph, as has been suggested, data from this study suggests that the selective basis for the establishment of floral polymorphism could have been increased pollen transfer rather than higher levels of outcrossing.     

THE INTERPLAY BETWEEN LOCAL ECOLOGY,DIVERGENT SELECTION,AND GENETIC DRIFT IN POPULATION DIVERGENCE OF A SEXUALLY ANTAGONISTIC FEMALE TRAIT

Genetically polymorphic species offer the possibility to study maintenance of genetic variation and the potential role for genetic drift in population divergence. Indirect inference of the selection regimes operating on polymorphic traits can be achieved by comparing population divergence in neutral genetic markers with population divergence in trait frequencies. Such an approach could further be combined with ecological data to better understand agents of selection. Here, we infer the selective regimes acting on a polymorphic mating trait in an insect group; the dorsal structures (either rough or smooth) of female diving beetles. Our recent work suggests that the rough structures have a sexually antagonistic function in reducing male mating attempts. For two species (Dytiscus lapponicus and Graphoderus zonatus), we could not reject genetic drift as an explanation for population divergence in morph frequencies, whereas for the third (Hygrotus impressopunctatus) we found that divergent selection pulls morph frequencies apart across populations. Furthermore, population morph frequencies in H. impressopunctatus were significantly related to local bioclimatic factors, providing an additional line of evidence for local adaptation in this species. These data, therefore, suggest that local ecological factors and sexual conflict interact over larger spatial scales to shape population divergence in the polymorphism.