EVOLUTION OF TRADE-OFFS BETWEEN SEXUAL AND ASEXUAL PHASES AND THE ROLE OF REPRODUCTIVE PLASTICITY IN THE GENETIC ARCHITECTURE OF APHID LIFE HISTORIES

Life‐history theory postulates that evolution is constrained by trade‐offs (i.e., negative genetic correlations) among traits that contribute to fitness. However, in organisms with complex life cycles, trade‐offs may drastically differ between phases, putatively leading to different evolutionary trajectories. Here, we tested this possibility by examining changes in life‐history traits in an aphid species that alternates asexual and sexual reproduction in its life cycle. The quantitative genetics of reproductive and dispersal traits was studied in 23 lineages (genotypes) of the bird cherry‐oat aphid Rhopalosiphum padi, during both the sexual and asexual phases, which were induced experimentally under specific environmental conditions. We found large and significant heritabilities (broad‐sense) for all traits and several negative genetic correlations between traits (trade‐offs), which are related to reproduction (i.e., numbers of the various sexual or asexual morphs) or dispersal (i.e., numbers of winged or wingless morphs). These results suggest that R. padi exhibits lineage specialization both in reproductive and dispersal strategies. In addition, we found important differences in the structure of genetic variance–covariance matrices ( G ) between phases. These differences were due to two large, negative genetic correlations detected during the asexual phase only: (1) between fecundity and age at maturity and (2) between the production of wingless and winged parthenogenetic females. We propose that this differential expression in genetic architecture results from a reallocation scheme during the asexual phase, when sexual morphs are not produced. We also found significant G × E interaction and nonsignificant genetic correlations across phases, indicating that genotypes could respond independently to selection in each phase. Our results reveal a rather unique situation in which the same population and even the same genotypes express different genetic (co)variation under different environmental conditions, driven by optimal resource allocation criteria.     

Can Coral Cover be Restored in the Absence of Natural Recruitment and Reef Recovery?

With many coral reef areas being degraded whether by anthropogenic or natural causes, a search is on for resilient species of corals that can restore coral cover where needed, if coral reefs are to continue to provide adequate ecosystem services. A series of experiments were undertaken in two sites with different environmental attributes and substrates in a lagoonal area in the northwestern Philippines to test the potentials of a local species, Porites cylindrica, for reef rehabilitation. With the use of asexual fragmentation of donor colonies, different treatments were tested, particularly to determine if the species would survive on different substrates, that is, solid, massive versus digitate/anastomosing, dead colonies. The results after nearly 2 years of the experiment were extraordinarily successful, with survival of transplants ranging from a high of 98% to a low of 80% of colonies, resulting in extensive coral cover on both original and new or different substrate from the original. A subsequent observation after another 16 months showed the coral cover to have been complete or nearly complete in the experimental plots, with the transplanted colonies fusing, and with evident reef fish communities where there were none before. Had there been no intervention, it is highly likely that the reefs would have remained in a degraded state .     

The effects of sexual and asexual reproduction on geographic variation in the sea anemone Actinia tenebrosa

Allelic and genotypic frequencies were determined for samples from 35 widely distributed Australasian colonies of Actinia tenebrosa and 2 South African colonies of A. equina. These data provided no evidence of gene flow between Australisian and South African Actinia colonies and indicated that there may be some restriction of gene flow between widely separated Australasian colonies.Both species are viviparous, and brooded A. tenebrosa are known to be produced asexually. The present data indicate that, within both species, almost all genotypic diversity is generated by sexual reproduction with recombination. Sexually produced juveniles appear to be widely dispersed and panmixis may occur over thousands of kilometres. However, successful sexual recruitment must be episodic or rare. Colonies on stable shores displayed relatively low levels of genotypic diversity, as compared with expectations for sexually reproducing populations, indicating strong local effects of asexual recruitment. Clonal genotypes may be spread over hundreds of metres of shore, but are typically restricted to discrete colonies. Asexual recruitment is highly localised and asexual dispersal appears to be limited by lengths of shore (500 m) which are unsuitable for colonization. Colonies on unstable shores are significantly more diverse genotypically and show little evidence of clonal proliferation.     

Recruitment failure in Florida Keys <Emphasis Type="Italic">Acropora palmata,</Emphasis> a threatened Caribbean coral

Recovery of Acropora palmata from its currently imperiled status depends on recruitment, a process which is poorly documented in existing Caribbean coral population studies. A. palmata is thought to be well adapted to proliferate through the recruitment of fragments resulting from physical disturbances, such as moderate intensity hurricanes. This study monitored fifteen 150 m² fixed study plots on the upper Florida Keys fore-reef for asexual and sexual recruitment from 2004 to 2007. Between July and October 2005, 4 hurricanes passed by the Florida Keys, producing wind speeds on the reef tract of 23 to 33 m s⁻¹. Surveys following the hurricanes documented an average loss of 52% estimated live tissue area within the study plots. The percentage of “branching” colonies in the population decreased from 67% to 42% while “remnant” colonies (isolated patches of tissue on standing skeleton) increased from 11% to 27%. Although some detached branches remained as loose fragments, more than 70% of the 380 fragments observed in the study plots were dead or rapidly losing tissue 3 weeks after Hurricane Dennis. Over the course of the study, only 27 fragments became attached to the substrate to form successful asexual recruits. Meanwhile, of the 18 new, small encrusting colonies that were observed in the study, only 2 were not attributable to asexual origin (i.e., remnant tissue from colonies or fragments previously observed) and are therefore possible sexual recruits. In summary, the 2005 hurricane season resulted in substantial loss of A. palmata from the upper Florida Keys fore-reef from a combination of physical removal and subsequent disease-like tissue mortality, and yielded few recruits of either sexual or asexual origin. Furthermore, the asexual and sexual fecundity of the remaining population is compromised for the near future due to the lack of branches (i.e., “asexual fecundity”) and overall loss of live tissue.     

Parasite resistance predicts fitness better than fecundity in a natural population of the freshwater snail Potamopyrgus antipodarum

The cost of males should give asexual females an advantage when in competition with sexual females. In addition, high‐fecundity asexual genotypes should have an advantage over low‐fecundity clones, leading to reduction in clonal diversity over time. To evaluate fitness components in a natural population, we measured the annual reproductive rate of individual sexual and asexual female Potamopyrgus antipodarum, a New Zealand freshwater snail, in field enclosures that excluded competitors and predators. We used allozyme genotyping to assign the asexual females to particular clonal genotypes. We found that the most fecund asexual clones had similar or higher fecundity as the top 10% of sexual families, suggesting that fecundity selection, even without the cost of males, would lead to replacement of the sexual population by clones. Consequently, we expected that the clones with the highest fecundity would dominate the natural population. Counter to this prediction, we found that high annual reproductive rates did not correlate with the frequency of clones in the natural population. When we exposed the same clones to parasites in the laboratory, we found that resistance to infection was positively correlated with the frequency of clones in the population. The correlation between fecundity and parasite resistance was negative, suggesting a trade‐off between these two traits. Our results thus suggest that parasite resistance is an important short‐term predictor of the success of asexual P. antipodarum in this population.     

An experimental analysis of pleometrotic advantage in the desert seed-harvester antMessor pergandei (Hymenoptera; Formicidae)

Summary Starting colonies of the desert seed-harvester antMessor pergandei are clumped in the field and face severe intraspecific competition through brood raiding. Single foundress laboratory colonies ofM. pergandei are more likely to succeed at brood raiding with conspecific colonies if they are given additional workers and mature pupae several days prior to brood raiding. Per foundress fecundity remains constant across laboratory starting colonies established with 1, 3 and 5 foundresses. These results suggest that the selective advantage of cooperative colony foundation (pleometrosis) in this and similar species may derive directly from the ability of multiple foundresses to produce a larger brood raiding force.     

THE LIFE CYCLE OF PEDIASTRUM SIMPLEX1

A sexual cycle and a newly discovered mode of asexual reproduction by azygotes arc described for Pediastrum simplex. In the sexual cycle isogametes from colony cells fuse in pairs, develop into zygotes, which grow and divide, producing motile cells which become angular polyeders. Polyeders may divide internally to form 4-celled colonies, or they enlarge, cleave, and produce zoospores or nonmotile cells. Zoospores are released in vesicles, swarm, adhere to each other, and form, flat colonies I cell thick. The nonmotile cells adhere to each other, enlarge, break out of the polyeder, and become spheroidal colonies. In the asexual cycle, nonfusing gametes, and motile cells derived from colony cells, become 4-spined azygotes which enlarge, divide internally, and release motile cells developing into polyeders. These polyeders develop further like those from zygotes.     

ORIGIN AND MAINTENANCE OF A FEMALE MATING PREFERENCE

We investigated the role of maintenance and origin of female mating preferences in three species of fish. Poecilia latipinna and P. mexicana are sexual species that hybridized to form the gynogenetic clonal P. formosa, which reproduces asexually but requires sperm to initiate embryogenesis. We demonstrate that all three species display almost identical and statistically indistinguishable preferences for large males. Although processes of good genes, runaway sexual selection, and direct selection could maintain preferences in the sexual species, good genes and runaway sexual selection are unlikely to operate in the asexual species. Furthermore, we found that the most likely direct selection benefit, an increase in fecundity, can also be excluded in the gynogens. We conclude that the most parsimonious explanation for this P. formosa preference is that it was inherited from the parental species and is maintained without forces generated by good genes, runaway selection, or direct selection for increased fecundity. This preference may be maintained because of pleiotropic effects (e.g., sensory bias) or mate searching costs.     

Evolutionary implications of polygyny in the Argentine ant,Iridomyrmex humilis (Mayr) (Hymenoptera: Formicidae): an experimental study

A comparison of several physiological parameters of queens of Iridomyrmex humilis in experimental monogynous and polygynous colonies showed that queens in monogynous colonies became heavier, had more developed ovaries and laid about twice as many eggs. Workers in monogynous colonies were more attracted to queens, which therefore probably received more food. This may partially explain the higher weight and fecundity of queens in monogynous colonies of Iridomyrmex humilis and possibly other ant species. In polygynous colonies, queens differed greatly in their fecundity. These differences did not appear to be the result of a dominance hierarchy. These results are discussed from an evolutionary point of view. Two hypotheses of mutualism and colony level selection are proposed as an alternative to kin selection which is unlikely to be the exclusive selective influence in the evolution of polygyny either in I. humilis or in most other ant species.     

The Effect of Queen and Worker Adoption on Weaver Ant (Oecophylla smaragdina F.) Queen Fecundity

Incipient ant colonies are often under fierce competition, making fast growth crucial for survival. To increase production, colonies can adopt multiple queens (pleometrosis), fuse with other colonies or rob brood from neighboring colonies. However, different adoption strategies might have different impacts such as future queen fecundity or future colony size. O. smaragdina queen production was measured in incipient colonies with 2, 3 or 4 founding queens, following the transplantation of 0, 30 or 60 pupae from a donor colony. Pupae developed into mature workers, resulting in increased worker/queen ratios in pupae transplanted treatments and leading to increases in the per capita queen production. Conversely, more queens did not induce increased per capita fecundity. Thus, brood robbing added individuals to the worker force and increased future production of resident queens, whereas queen adoption increased the colony’s future production, but not the production of individual queens.     

POPULATION BIOLOGY OF AVENA. VIII. COLONIZATION EXPERIMENT AS A TEST OF THE ROLE OF NATURAL SELECTION IN POPULATION DIVERGENCE

Artificially founded colonies of Avena barbata were utilized in two experiments in order to examine the role of natural selection in the evolution of regional differentiation in Central California. One experiment involved a total of 41 colonies founded in three different areas and scored over a 10-yr period; these colonies, started with known genotypes, showed that although both Valley and Bay region genotypes establish successful colonies, their relative survivorship and fecundity suggests weak selective forces consistent with those predicted from the observed patterns in natural populations. The second experiment involved two localities, representing two climatic regimes of temperature, in each of which ten colonies were started from identical seed sources. These colonies also showed relatively higher fitness of the genotypes matching with those sampled from the Valley and Coastal regions, respectively, again in line with their regional pattern of distribution. Colonization experiments appeared to be useful even as short-term in situ tests of certain specific evolutionary hypotheses on the role and intensity of natural selection.     

Changes in gametogenesis and fecundity of acroporid corals that were exposed to elevated nitrogen and phosphorus during the ENCORE experiment

Colonies of two scleractinian reef coral species, Acropora longicyathus and Acropora aspera were transplanted into patch reefs at One Tree Reef, Great Barrier Reef, Australia as part of the ENCORE experiment. These corals and colonies of A. aspera which were naturally present in the patch reefs were exposed to four treatments over two years: controls with normal seawater, elevated levels of nitrogen only, phosphorus only, or nitrogen plus phosphorus. These corals were sampled and used to determine whether gametogenic cycles and fecundity were affected by nutrient enrichment. Acropora longicyathus had a single annual gametogenic cycle. Corals exposed to elevated nitrogen produced significantly smaller and fewer eggs and contained less testes material than those which were not exposed to nitrogen. Exposure to elevated phosphorus only resulted in corals producing more but smaller eggs, and more testes material. Egg numbers of colonies from other treatments decreased as the gametogenic cycles continued, but those of the phosphorus colonies showed almost no reduction in egg numbers between the early and late stages of the gametogenic cycles. These results have important management implications for coral reefs as they demonstrate that small increases in concentrations of nitrogen and phosphorus can have severe effects on reproductive activity in these species of scleractinian corals.     

Enhanced ultraviolet radiation can terminate sexual reproduction in the broadcasting coral species Acropora cervicornis Lamarck

The effects of enhanced ultraviolet radiation (280-400 nm: UVR) on the fecundity of Acropora cervicornis were measured in field-transplanted colonies from 20 m to 1 m depth and vice versa at La Parguera, Puerto Rico. Fecundity was estimated from histological sections made from tissue samples obtained at different time intervals during the experimental period (March - August 2003). All colonies transplanted from 20 m to 1 m showed a 100% reduction in gonads per mesentery per polyp one month after transplantation, while those transplanted from 1 m to 20 m did not show any significant reduction in fecundity throughout the experimental period. The latter colonies did show however, a delay in the spawning times by releasing their gamete bundles approximately two-three weeks after the controls at 1 m and two months after the controls at 20 m suggesting an induced response as a consequence of changes in their daily light cycle due to less radiation (PAR and UVR) available at 20 m compared to 1 m. Control colonies at 20 m spawned after the full moon of June 2003, while the controls at 1 m spawned 5-6 days after the full moon of July 2003. While a possible reabsorption of the gametes occurred in A. cervicornis colonies transplanted from 20 m to 1 m, the expulsion of these gametes due to the sudden stress caused by the transplantation is not discarded. The results suggest that sudden increases in UVR can completely stop sexual reproduction in ramose broadcasting coral species, which in turn can affect the dominance of the species and the composition and structure of shallow reef environments.     

EVOLUTION OF ASEXUALITY VIA DIFFERENT MECHANISMS IN GRASS THRIPS (THYSANOPTERA: Aptinothrips)

Asexual lineages can derive from sexual ancestors via different mechanisms and at variable rates, which affects the diversity of the asexual population and thereby its ecological success. We investigated the variation and evolution of reproductive systems in Aptinothrips, a genus of grass thrips comprising four species. Extensive population surveys and breeding experiments indicated sexual reproduction in A. elegans, asexuality in A. stylifer and A. karnyi, and both sexual and asexual lineages in A. rufus. Asexuality in A. stylifer and A. rufus coincides with a worldwide distribution, with sexual A. rufus lineages confined to a limited area. Inference of molecular phylogenies and antibiotic treatment revealed different causes of asexuality in different species. Asexuality in A. stylifer and A. karnyi has most likely genetic causes, while it is induced by endosymbionts in A. rufus. Endosymbiont‐community characterization revealed presence of Wolbachia, and lack of other bacteria known to manipulate host reproduction. However, only 69% asexual A. rufus females are Wolbachia‐infected, indicating that either an undescribed endosymbiont causes asexuality in this species or that Wolbachia was lost in several lineages that remained asexual. These results open new perspectives for studies on the maintenance of mixed sexual and asexual reproduction in natural populations.     

The influence of prior experience on preference and performance of a cryptoparasitoid Scleroderma guani (Hymenoptera: Bethylidae) on beetle hosts

1. Numerous studies have reported the effects of learning or experience on parasitoid host preference and location. However, the integration of pre‐imaginal and adult experiences on the subsequent host preference and adult/offspring performance has been rarely tested in host–parasite interactions. 2. We present direct evidence that theses two kinds of experiences affect host preference and related fitness in the polyphagous parasitoid, Scleroderma guani. Two colonies of parasitoids were reared on Monochamus alternatus and Saperda populnea (Cerambycidae: Lamiinae). Individuals from the two colonies were given host‐switching experience for one generation (pre‐imaginal experience) while other individuals were given prior ovipositing experience on the two species, respectively (adult experience). 3. Scleroderma guani females demonstrated that their experiences determined adult behavioural responses and their subsequent performance to hosts. Females maximised both adult fitness (fecundity and longevity) and offspring fitness (survival and sex ratio) when they encountered hosts similar to their maternal hosts. Behavioural plasticity in host choice was affected by adult experience, resulting in improved adult feeding and ovipositing behaviour and further modifying adult fecundity and the offspring sex ratio. There was a positive correlation between oviposition preference and adult fecundity. 4. The results indicated that S. guani exhibited positive preference–performance correlations. This is most likely due to an adaptation to maternal hosts over multiple generations. However, foraging potential of adults to available cues from hosts may be driven quickly by an experience‐induced learning process rather than by natural selection processes shaped over many generations.     

POPULATION PERSISTENCE AND REPRODUCTIVE ECOLOGY OF A FOREST HERB: ASTER ACUMINATUS

Aster acuminatus Michaux, a patchily distributed herb of eastern forests, is commonly represented in all stages of forest succession. To determine its mechanisms of persistence, we carefully examined the distribution patterns of this species as affected by its reproductive ecology and demography. Using A. acuminatus as a model, we hypothesized that 1) availability of propagules and location of adequate recruitment sites impose first-order control over plant distributions in stands of all ages by determining where plants might be found and 2) light and other resources exert a second-order control over herb distributions by determining where established plants cannot persist. Results from our study at the Hubbard Brook Experimental Forest in New Hampshire demonstrated this species' ability to exploit resource-rich disturbances through immediate and profuse ramet and seed production. This ensured rapid colonization of new sites and provided genetic exchange for longer-term survival. Relict colonies which survived the disturbance served as the sole propagule-producing loci for the first year of colonization. After several years, resources became more limiting and the aster shifted to a more conservative asexual reproduction strategy which provided population stability until the next disturbance. The clumped distribution of A. acuminatus in mature forests is explained by strict adherence to asexual ramet replacement in which there is no year-to-year change in aster importance. Location of persistent colonies probably is dictated by location of suitable seedbeds during the first few years of recovery after disturbance. Whereas numerous sites may be colonized initially, timing of colonization and quality of site impose constraints on which colonies persist.     

Developmental changes in factors limiting colony survival and growth of the leaf‐cutter ant Atta laevigata

Many species of leaf‐cutter ants (Atta and Acromyrmex) increase in abundance following natural or anthropogenic disturbances in the vegetation. However, the mechanisms responsible for such an increase are still poorly understood. We evaluated the effects of nesting site and the availability of palatable forage on survival and growth of Atta laevigata colonies at different developmental stages. Foundress queens transplanted into man‐made clearings (dirt roads) had a much higher survival than those transplanted into the adjacent undisturbed savannah vegetation. Similarly, incipient colonies (≥3‐months old) had significantly greater survival and growth rates in dirt roads. In contrast, nesting site did not influence performance of young colonies (≥15‐months old). Both incipient and young colonies responded strongly and positively to experimental supplementation of palatable forage, and this effect was independent of the nesting habitat. Colonies that received extra food grew faster and had a significantly greater survival rate than control colonies. These results suggest that performance of A. laevigata is affected by the generally greater availability of suitable nesting sites and palatable vegetation in disturbed habitats. This may explain how these ants maintain high densities in these habitats, and since the relative importance of these factors changed with colony ontogeny, our findings highlight the importance of evaluating potential limiting factors throughout the full range of an organism's developmental stages.     

Adaptation in the Asexual False Spider Mite <Emphasis Type="Italic">Brevipalpus phoenicis</Emphasis>: Evidence for Frozen Niche Variation

Because asexual species lack recombination, they have little opportunity to produce genetically variable offspring and cannot adapt to changes in their environment. However, a number of asexual species are very successful and appear to contradict this general view. One such species is the phytophagous mite Brevipalpus phoenicis (Geijskes), a species that is found in a wide range of environments. There are two general explanations for this pattern, the General Purpose Genotype (GPG) and Frozen Niche Variation (FNV). According to the GPG model, an asexual species consists of clones that can all survive and reproduce in all the different niches. Alternatively, the FNV model postulates that different clones are specialized to different niches. We have performed a test to distinguish between these models in B. phoenicis. Mites from three populations from three different host plant species (citrus, hibiscus and acerola) were transplanted to their own and the two alternative host plants and mite survival and egg production were measured. Additionally, the mite populations were genotyped using microsatellites. Fitness was seriously reduced when mites were transplanted to the alternative host plant species, except when the alternative host was acerola. We concluded that B. phoenicis clones are specialized to different niches and thus the FNV best describes the broad ecological niche of this species but that there is also some evidence for host plant generalization. This conclusion was strengthened by the observations that on each host plant species the native mite population performed better than the introduced ones, and that three microsatellite markers showed that the mite populations are genetically distinct.     

Equal fecundity in asexual and sexual mollies (<Emphasis Type="Italic">Poecilia</Emphasis>)

The evolution and maintenance of sexual reproduction is still one of the major unresolved problems in evolutionary biology. Sexual reproduction is fraught with a number of costs as compared to asexual reproduction. For example, sexuals have to produce males, which–given a 1:1 sex ratio—results in a two-fold advantage for asexuals that do not produce males. Consequently, asexuals will outperform and replace sexuals over time assuming everything else is equal. Nonetheless, a few cases of closely related asexuals and sexuals have been documented to coexist stably in natural systems. We investigated the presence of a two-fold cost in a unique system of three closely related fish species: the asexual Amazon Molly (Poecilia formosa), and two sexual species, Sailfin Molly (P. latipinna) and Atlantic Molly (P. mexicana). Amazon Molly reproduce gynogenetically (by sperm dependent parthenogenesis) and always coexist with one of the sexual species, which serves as sperm donor. In the laboratory, we compared reproductive output between P. formosa and P. mexicana as well as P. formosa and P. latipinna. We found no differences in the fecundity in either comparison of a sexual and the asexual species. Under the assumption of a 1:1 sex ratio, the asexual Amazon Molly should consequently have a full two-fold advantage and be able to outcompete sexuals over time. Hence, the coexistence of the species pairs in nature presents a paradox still to be solved.     

ASEXUAL AND SEXUAL REPRODUCTION IN GONIUM QUADRATUM (CHLOROPHYTA) WITH A DISCUSSION OF PHYLOGENETIC RELATIONSHIP WITHIN THE GONIACEAE1

Morphological details of asexual and sexual reproduction in Gonium Quadratum Pringsheim ex Nozaki (Goniaceae, Chlorophyta) were observed by light microscopy, based on clonal cultured materials originating from Nepal. In asexual reproduction, the alga exhibited two different patterns of cell cleavage during formation of 8-and 16-celled daughter colonies. Sexual reproduction was heterothallic and isogamous. The gametes bore a tubular mating structure (bilateral mating papilla) at the base of the flagella, and the papillae of the two gametes. The germinating zygote gave rise to four biflagellate gone cells joined in a colony (germ colony). Possible phylogenetic relationships within the Goniaceae at the species level are outlined, mainly on the basis of reproduction characteristics.