Asexual Origin Research Articles

The role of hybridization in the origin and spread of asexuality in Daphnia

The molecular mechanisms leading to asexuality remain little understood despite their substantial bearing on why sexual reproduction is dominant in nature. Here, we examine the role of hybridization in the origin and spread of obligate asexuality in Daphnia pulex, arguably the best-documented case of contagious asexuality. Obligately parthenogenetic (OP) clones of D.pulex have traditionally been separated into 'hybrid' (Ldh SF) and 'nonhybrid' (Ldh SS) forms because the lactase dehydrogenase (Ldh) locus distinguishes the cyclically parthenogenetic (CP) lake dwelling Daphnia pulicaria (Ldh FF) from its ephemeral pond dwelling sister species D.pulex (Ldh SS). The results of our population genetic analyses based on microsatellite loci suggest that both Ldh SS and SF OP individuals can originate from the crossing of CP female F1 (D.pulex×D.pulicaria) and backcross with males from OP lineages carrying genes that suppress meiosis specifically in female offspring. In previous studies, a suite of diagnostic markers was found to be associated with OP in Ldh SS D.pulex lineages. Our association mapping supports a similar genetic mechanism for the spread of obligate parthenogenesis in Ldh SF OP individuals. Interestingly, our study shows that CP D.pulicaria carry many of the diagnostic microsatellite alleles associated with obligate parthenogenesis. We argue that the assemblage of mutations that suppress meiosis and underlie obligate parthenogenesis in D.pulex originated due to a unique historical hybridization and introgression event between D.pulex and D.pulicaria.

Microsatellite and single‐nucleotide polymorphisms indicate recurrent transitions to asexuality in a microsporidian parasite

Assessing the mode of reproduction of microparasites remains a difficult task because direct evidence for sexual processes is often absent and the biological covariates of sex and asex are poorly known. Species with geographically divergent modes of reproduction offer the possibility to explore some of these covariates, for example, the influence of life-history traits, mode of transmission and life-cycle complexity. Here, we present a phylogeographical study of a microsporidian parasite, which allows us to relate population genetic structure and mode of reproduction to its geographically diverged life histories. We show that in microsporidians from the genus Hamiltosporidium, that use the cladoceran Daphnia as host, an epidemic population structure has evolved, most probably since the last Ice Age. We partially sequenced three housekeeping genes (alpha tubulin, beta tubulin and hsp70) and genotyped seven microsatellite loci in 51 Hamiltosporidium isolates sampled within Europe and the Middle East. We found two phylogenetically related asexual parasite lines, one each from Fennoscandia and Israel, which share the unique ability of being transmitted both vertically and horizontally from Daphnia to Daphnia. The sexual forms cannot transmit horizontally among Daphnia, but presumably have a complex life cycle with a second host species. In spite of the similarities between the two asexual lineages, a clustering analysis based on microsatellite polymorphisms shows that asexual Fennoscandian parasites do not share ancestry with any other Hamiltosporidium that we have sampled. Moreover, allele sequence divergence at the hsp70 locus is twice as large in Fennoscandian than in Israeli parasites. Our results indicate that asexual reproduction evolved twice independently, first in Fennoscandian and more recently in the Israeli parasites. We conclude that the independent origin of asexuality in these two populations is associated with the altered parasite mode of transmission and the underlying dynamics of host populations.

Dynamic Formation of Asexual Diploid and Polyploid Lineages: Multilocus Analysis of Cobitis Reveals the Mechanisms Maintaining the Diversity of Clones

Given the hybrid genomic constitutions and increased ploidy of many asexual animals, the identification of processes governing the origin and maintenance of clonal diversity provides useful information about the evolutionary consequences of interspecific hybridization, asexuality and polyploidy. In order to understand the processes driving observed diversity of biotypes and clones in the Cobitis taenia hybrid complex, we performed fine-scale genetic analysis of Central European hybrid zone between two sexual species using microsatellite genotyping and mtDNA sequencing. We found that the hybrid zone is populated by an assemblage of clonally (gynogenetically) reproducing di-, tri- and tetraploid hybrid lineages and that successful clones, which are able of spatial expansion, recruit from two ploidy levels, i.e. diploid and triploid. We further compared the distribution of observed estimates of clonal ages to theoretical distributions simulated under various assumptions and showed that new clones are most likely continuously recruited from ancestral populations. This suggests that the clonal diversity is maintained by dynamic equilibrium between origination and extinction of clonal lineages. On the other hand, an interclonal selection is implied by nonrandom spatial distribution of individual clones with respect to the coexisting sexual species. Importantly, there was no evidence for sexually reproducing hybrids or clonally reproducing non-hybrid forms. Together with previous successful laboratory synthesis of clonal Cobitis hybrids, our data thus provide the most compelling evidence that 1) the origin of asexuality is causally linked to interspecific hybridization; 2) successful establishment of clones is not restricted to one specific ploidy level and 3) the initiation of clonality and polyploidy may be dynamic and continuous in asexual complexes.

Unusual pattern of embryogenesis of Caryophyllia inornata (scleractinia, caryophylliidae) in the mediterranean sea: Maybe agamic reproduction?

While knowledge of the reproductive biology of tropical scleractinian corals is extensive, information from temperate zones is limited. The aim of this study is to describe the reproductive biology of Caryophyllia inornata, a temperate species, and to increase the understanding of the reproductive strategies of Mediterranean corals. Samples of C. inornata were collected during SCUBA surveys at Elba island. Sexually active individuals displayed either male or female germ cells, showing a gonochoric sexuality. C. inornata exhibited an unusual pattern of embryogenesis. Embryos appeared throughout the whole year in males and in sexually inactive individuals, and they did not show a seasonal pattern of development, as usually expected for sexual reproduction. This observation suggests the possibility of asexual origin. These embryogenetic sexually inactive individuals were larger in size than the embryogenetic sexually active ones, and they might be senile polyps that preserve the ability to produce embryos only by agamic reproduction.

Differential effects of polyploidy and diploidy on fitness of apomictic Boechera

The co-occurrence of apomixis (asexual reproduction) and polyploidy in plants has been the subject of debate in regard to the origin and evolution of asexuality. In recent years, polyploidy has been postulated as a maintenance and stabilization factor rather than as a source of apomixis origin. It is assumed polyploidy facilitates the compensation for mutation accumulation, and hence, the rare occurrence of diploid apomixis indirectly supports this finding. Nevertheless, diploid apomicts exist and are successful, especially in the genus Boechera. While comparing phenotypic traits, fitness-related traits and apomixis penetrance between both diploid and triploid apomicts in the genus Boechera, it was expected to find trait variance that can be attributed to ploidy. Surprisingly, little trait variation could be assigned to ploidy, but rather trait variations were mainly genotype-specific. Additionally, it is shown that paternal contribution is very important for trait success, even though all offspring are genetically identical to the mother plant. This harbors implications for the introduction of apomixis into crop plants, considering the effects of paternal contribution during asexual reproduction. Nevertheless, polyploidy is an efficient way to buffer deleterious mutations, but the flexibility of diploid apomicts of the genus Boechera for rare sexual events contributes to their success in nature.

Evolution and biogeography of alpine apomictic plants

AbstractAsexual organisms often occupy larger and more northern distribution areas than their sexual relatives, and tend to colonize more frequently previously glaciated areas. These phenomena, summarized under the term "geographical partheno ‐ genesis", seem to imply a short‐term advantage of asexual reproduction. Alpine biota, for instance, with their short growing seasons, cold climates and climatic histories shaped by glaciations are predisposed for the evolution and spread of apomictic plants. However, patterns and causes of geographical parthenogenesis for alpine species remain elusive, and for many alpine taxa the mode of reproduction is only poorly characterized. A survey of apomictic species in the European Alps presents some examples and case studies on geographical parthenogenesis. The distribution pattern may be explained by better colonizing abilities of asexual organisms because of uniparental reproduction. Apomixis has been advantageous for re‐colonization of the Alps after the retreat of glaciers, because of their ability to rapidly found populations via single individuals (Baker's Law). Asexual organisms also may perform better in diverse and narrow ecological niches. The distributional superiority of asexuals has also been attributed to indirect advantages of hybridity and/or polyploidy. Sexual hybrids or polyploids, however, when compared to sexual diploid progenitors, do not show patterns of geographical parthenogenesis. However, in the Alps, climatic oscillations may have triggered range fluctuations of species and breakdown of crossing barriers. Interspecific hybridization and polyploidy may have spurred recurrent origins of asexuality, while decreasing the fitness of sexual progenitor species. Biogeographical history in combination with the intrinsic advantages of apomixis may explain best the relative success of alpine apomictic plants.

Tropical UnderstoryPiperShrubs Maintain High Levels of Genotypic Diversity Despite Frequent Asexual Recruitment

ABSTRACTMany plant species have the capacity to regenerate asexually by resprouting from stem and leaf fragments. In the pan‐tropical shrub genusPiper, this tendency is thought to be higher in shade‐tolerant than light‐demanding species, and to represent a trade‐off with annual seed production. Here we use molecular markers to identify clones in fivePiperspecies varying in light requirements. We test predictions that (i) asexual recruitment success is highest in shade‐tolerant species, and (ii) that consequently, shade‐tolerant species are characterized by lower genotypic diversity than light‐demandingPiper. We found that two shade‐tolerantPiperspecies recruited asexually more frequently (36–42% of sampled shoots were of asexual origin) than, two light‐demanding and one shade‐tolerant species (0–26%). Furthermore, as predicted, genotypic diversity was negatively correlated with the frequency of asexual recruitment in the population. Nonetheless, genotypic diversity ofPiperwas high compared with other clonal plants. The proportion of unique genotypes found per population ranged from 0.58 to 1.0 and the genotypic Simpson's diversity ranged from 0.93 to 1.0 for all five species. Our results suggest that even though asexual reproduction plays an important role in maintaining local populations ofPiperin the understory, it does not seem to reduce genotypic diversity to levels that will threaten these species ability to respond to environmental change.Abstract in Spanish is available in the online version of this article.

Frozen F1’s amidst a masterpiece of nature: new insights into the rare hybrid origin of gynogenesis in the Amazon molly (Poecilia formosa)

All-female 'species' of fish have been shown to be great models in ecological and evolutionary studies because of the insights they can provide into the origin and evolution of asexuality, the ecology of hybrids, associations between genotype and environment, and the maintenance of sex. Gynogenetic organisms that evolved from sexual ancestors, and combine the disadvantageous traits from sexuality and asexuality, have long baffled evolutionary biologists trying to understand their origin and persistence with their sympatric sexual counterparts. In this issue, a new study using an integrated molecular phylogenetic and classical genetic approach has uncovered compelling evidence regarding the obscure asexual origin of the Amazon molly, Poecilia formosa. By performing an extensive phylogeographic analysis, Stöck etal. (2010) provide evidence that the Amazon molly arose only once within its history, with monophyly being strongly supported by mitochondrial DNA and microsatellite analyses. This result, combined with an elaborate failed attempt to resynthesize the lineage, suggests that vertebrate gynogens such as the Amazon molly are not rare because they are at a disadvantage to their sexual counterparts, but because the genomic conditions under which they arise are rare. Organisms that apparently combine the disadvantages of both sexuality and asexuality remain difficult to understand from both an ecological and an evolutionary perspective, and Stöck etal. (2010) highlight several outstanding important questions. Nonetheless, given that we now have a better knowledge of the origin and history of this unique 'species', this should allow researchers to better understand how these frozen F1's can persist amidst the masterpiece of nature.

Evolutionary and functional insights into reproductive strategies of aphids

Aphids are among the few organisms capable of reproducing either sexually or asexually. This plasticity in reproductive mode is viewed as an adaptive response to cope with seasonal changes. Clonal reproduction occurs during the growing season allowing rapid population increase, while sexual reproduction occurs during late summer and leads to frost-resistant eggs that can survive winter conditions. This shift between these two extreme reproductive modes is achieved by using the same genotype, i.e. within the same genetic clone, and is triggered by photoperiodic changes perceived by the aphid brain or visual system. Advances have been made recently to depict genetic programs that relate to the regulation of reproductive modes in aphids. These studies have benefited from the rapid development of genomic and post-genomic resources obtained through the International Aphid Genomics Consortium. Here, we underline the importance of several candidate genes in the switch from clonal to sexual reproduction in aphids and whose roles await full validation. Besides reproductive mode variation expressed at the genotypic level, aphid species also frequently encompass lineages which have lost the sexual phase and hence the alternating clonal and sexual reproductive phases of the life cycle. This coexistence of sex and asexual reproduction within the same species raises questions on its evolutionary and ecological significance. We summarize the knowledge accumulated to date on the maintenance of sex as well as on the origin and evolution of asexuality in aphids. By combining functional genomics, genetic and ecological approaches on reproductive plasticity and polymorphism, we hope to obtain an integrative view of the evolutionary forces shaping aphid reproductive strategies, from gene to population and species levels.

Reproductive and recruitment traits as indicators of the invasive potential of Kalanchoe daigremontiana (Crassulaceae) and Stapelia gigantea (Apocynaceae) in a Neotropical arid zone

Populations of two ornamental exotic species, Kalanchoe daigremontiana and Stapelia gigantea, have been recently detected inside a protected area containing arid ecosystems in Venezuela. We indirectly tested their invasive potential by characterizing their reproductive biology and recruitment patterns and comparing our estimates against Baker's Law and reproductive profiles reported for invasive plants. K. daigremontiana is autogamous, produces >16,000 seeds per plant and also reproduces clonally. Despite low seed viability (17.9%) and germination rates (11.9%), seeds were present in the seed bank. Plantlets of asexual origin showed high survival (75–100%) compared to seedlings of sexual origin (10%). S. gigantea is self-incompatible, xenogamous and produces close to 1500 seeds per m 2 of plant tissue. Seed viability (77%) and germination rates (62%) were high, but this species is not represented in the seed bank. It has a vegetative growth. A combination of reproductive and recruitment attributes, which match those considered in Baker's Law and others reported for invasive plants, confer K. daigremontiana and S. gigantea the potential to invade Neotropical arid zones; the former mainly through selfing and production of numerous asexual plantlets, and the latter through an association with a locally abundant pollinator and production of wind dispersed seeds.

MULTIPLE DIRECT TRANSITIONS FROM SEXUAL REPRODUCTION TO APOMICTIC PARTHENOGENESIS INTIMEMASTICK INSECTS

Transitions from sexual reproduction to parthenogenesis may occur along multiple evolutionary pathways and involve various cytological mechanisms to produce diploid eggs. Here, we investigate routes to parthenogenesis in Timema stick insects, a genus comprising five obligate parthenogens. By combining information from microsatellites and karyotypes with a previously published mitochondrial phylogeny, we show that all five parthenogens likely evolved spontaneously from sexually reproducing species, and that the sexual ancestor of one of the five parthenogens was probably of hybrid origin. The complete maintenance of heterozygosity between generations in the five parthenogens strongly suggests that eggs are produced by apomixis. Virgin females of the sexual species were also able to produce parthenogenetic offspring, but these females produced eggs by automixis. High heterozygosity levels stemming from conserved ancestral alleles in the parthenogens suggest, however, that automixis has not generated the current parthenogenetic Timema lineages but that apomixis appeared abruptly in several sexual species. A direct transition from sexual reproduction to (at least functional) apomixis results in a relatively high level of allelic diversity and high efficiency for parthenogenesis. Because both of these traits should positively affect the demographic success of asexual lineages, spontaneous apomixis may have contributed to the origin and maintenance of asexuality in Timema.

Recruitment failure in Florida Keys Acropora palmata, 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.

Genetic diversity and polyploidy in the cosmopolitan asexual ostracod Cypris pubera

Freshwater ostracods show extensive breeding system variation, but both the cause of breeding system transitions, and the origins of genotypic diversity in asexuals, have yet to be convincingly explained. The study reports on the genotypic (based on allozyme electrophoresis) characteristics of Northern European populations of the globally distributed ostracod Cypris pubera. No male has ever been described for C. pubera, and none were found in this study. Genotypic profiles indicated deviations from Hardy-Weinberg (HW) equilibrium, confirming that C. pubera reproduces through ameiotic parthenogenesis, that is, the production of female offspring where the formation of ova is through a mitotic process. Cypris pubera seems to be comprised of diploid clones that are brown in colour and a series of polyploid clones that are green. The two morphs often occurred in sympatry, but the green morph was considerably more diverse than the brown. The factors which contribute to asexual diversity or elevated ploidy levels in other taxa, such as hybridization, do not appear to be important for C. pubera. Thus, this study has established a link between polyploidy and genetic diversity in C. pubera, but the origins of both asexuality and polyploidy remain unclear for this ecologically successful ostracod.

Ice age cloning – comparison of the Quaternary evolutionary histories of sexual and clonal forms of spiny loaches (Cobitis; Teleostei) using the analysis of mitochondrial DNA variation

Recent advances in population history reconstruction offered a powerful tool for comparisons of the abilities of sexual and clonal forms to respond to Quaternary climatic oscillations, ultimately leading to inferences about the advantages and disadvantages of a given mode of reproduction. We reconstructed the Quaternary historical biogeography of the sexual parental species and clonal hybrid lineages within the Europe-wide hybrid complex of Cobitis spiny loaches. Cobitis elongatoides and Cobitis taenia recolonizing Europe from separated refuges met in central Europe and the Pontic region giving rise to hybrid lineages during the Holocene. Cobitis elongatoides due to its long-term reproductive contact with the remaining parental species of the complex--C. tanaitica and C. spec.--gave rise to two clonal hybrid lineages probably during the last interglacial or even earlier, which survived the Würmian glaciation with C. elongatoides. These lineages followed C. elongatoides postglacial expansion and probably decreased its dispersal rate. Our data indicate the frequent origins of asexuality irrespective of the parental populations involved and the comparable dispersal potential of diploid and triploid lineages.

EVOLUTIONARY HISTORY OF CONTAGIOUS ASEXUALITY IN DAPHNIA PULEX

Asexual taxa are short-lived, suggesting that transitions to asexuality represent evolutionary dead-ends. However, with high rates of clonal origin and coexistence of asexuals and sexuals via selective asymmetries, asexuality may persist in the long term as a result of a dynamic equilibrium between clonal origin and extinction. Few such systems have been studied in detail. Here, we investigate the evolutionary history of asexual lineages of Daphnia pulex, which are derived from sexual relatives via the inheritance of a dominant female-limited meiosis-suppressing locus and inhabit ponds throughout northeastern North America (NA). Our extensive sampling and subsequent phylogenetic analysis using mitochondrial sequence data reveals a young and genetically diverse asexual assemblage, reflecting high rates of clonal origin due to the contagious nature of asexuality. Yet, asexuality is restricted to two phylogroups (B and C) with historical and/or present associations with northeastern NA and is absent from a northwestern phylogroup (A), supporting a recent northeastern origin of asexuality in this species. Furthermore, macrogeographic patterns of genetic variability indicate that phylogroups B and C recolonized northeastern NA from opposite directions, yet their presently overlapping geographic distributions are similarly divided into an eastern asexual and a western sexual region. We attribute these patterns to a recent contagious spread of asexuality from a northeastern source. If environment-mediated selective asymmetries play no significant role in determining the outcome of competitive interactions between sexuals and asexuals, regions of contact may be setting the stage for continued asexual conquests.

EVOLUTIONARY HISTORY OF CONTAGIOUS ASEXUALITY IN DAPHNIA PULEX

Asexual taxa are short-lived, suggesting that transitions to asexuality represent evolutionary dead-ends. However, with high rates of clonal origin and coexistence of asexuals and sexuals via selective asymmetries, asexuality may persist in the long term as a result of a dynamic equilibrium between clonal origin and extinction. Few such systems have been studied in detail. Here, we investigate the evolutionary history of asexual lineages of Daphnia pulex, which are derived from sexual relatives via the inheritance of a dominant female-limited meiosis-suppressing locus and inhabit ponds throughout northeastern North America (NA). Our extensive sampling and subsequent phylogenetic analysis using mitochondrial sequence data reveals a young and genetically diverse asexual assemblage, reflecting high rates of clonal origin due to the contagious nature of asexuality. Yet, asexuality is restricted to two phylogroups (B and C) with historical and/or present associations with northeastern NA and is absent from a northwestern phylogroup (A), supporting a recent northeastern origin of asexuality in this species. Furthermore, macrogeographic patterns of genetic variability indicate that phylogroups B and C recolonized northeastern NA from opposite directions, yet their presently overlapping geographic distributions are similarly divided into an eastern asexual and a western sexual region. We attribute these patterns to a recent contagious spread of asexuality from a northeastern source. If environment-mediated selective asymmetries play no significant role in determining the outcome of competitive interactions between sexuals and asexuals, regions of contact may be setting the stage for continued asexual conquests.

PHYLOGENETIC EVIDENCE FOR HYBRID ORIGINS OF ASEXUAL LINEAGES IN AN APHID SPECIES

Understanding the mode of origin of asexuality is central to ongoing debates concerning the evolution and maintenance of sexual reproduction in eukaryotes. This is because it has profound consequences for patterns of genetic diversity and ecological adaptability of asexual lineages, hence on the outcome of competition with sexual relatives both in short and longer terms. Among the possible routes to asexuality, hybridization is a very common mechanism in animals and plants. Aphids present frequent transitions from their ancestral reproductive mode (cyclical parthenogenesis) to permanent asexuality, but the mode of origin of asexual lineages is generally not known because it has never been thoroughly investigated with appropriate molecular tools. Rhopalosiphum padi is an aphid species with coexisting sexual (cyclically parthenogenetic) and asexual (obligately parthenogenetic) lineages that are genetically distinct. Previous studies have shown that asexual lineages of R. padi are heterozygous at most nuclear loci, suggesting either that they have undergone long-term asexuality (under which heterozygosity tends to increase) or that they have hybrid origins. To discriminate between these alternatives, we conducted an extensive molecular survey combining the sequence analysis of alleles of two nuclear DNA markers and mitochondrial DNA haplotypes in sexual and asexual lineages of R. padi. Both nuclear and cytoplasmic markers clearly showed that many asexual lineages have hybrid origins, the first such demonstration in aphids. Our results also indicated that asexuals result from multiple events of hybridization between R. padi and an unknown sibling species, and are of recent origin (contradicting previous estimates that asexual R. padi lineages were of moderate longevity). This study constitutes another example that putatively ancient asexual lineages are actually of much more recent origin than previously thought. It also presents a robust approach for testing whether hybrid origin of asexuality is also a common phenomenon in aphids.

Multiple routes to asexuality in an aphid species.

Cyclical parthenogens, including aphids, are important models for studying the evolution of sex. However, little is known about transitions to asexuality in aphids, although the mode of origin of asexual lineages has important consequences for their level of genetic diversity, ecological adaptability and the outcome of competition with their sexual relatives. Thus, we surveyed nuclear, mitochondrial and biological data obtained on cyclical and obligate parthenogens of the bird cherry-oat aphid, Rhopalosiphum padi (L), to investigate the frequency of transitions from sexuality to permanent asexuality. Many instances of asexual lineages retaining the ability to produce males are known in aphids, so particular attention was paid to the existence of occasional matings between females from sexual lineages and males produced by asexual lineages, which have the potential to produce new asexual lineages. Phylogenetic inference based on microsatellite and mitochondrial data indicates at least three independent origins of asexuality in R. padi, yielding the strongest evidence to date for multiple origins of asexuality in an aphid. Moreover, several lines of evidence demonstrate that transitions to asexuality result from two mechanisms: a complete spontaneous loss of sex and repeated gene flow from essentially asexual lineages into sexual ones.

Reproductive patterns of Caloglossa species (Delesseriaceae, Rhodophyta) from Australia and New Zealand: multiple origins of asexuality in C. leprieurii. Literature review on apomixis, mixed-phase, bisexuality and sexual compatibility

SUMMARY Reproduction and life history patterns in culture of five Caloglossa speaes from Australia and New Zealand are compared. Caloglossa adhaerens King et Puttock and Caloglossa bengalensis (Martens) King et Puttock have a Polyslphonla-type sexual life history (P-type, isomorphic alternation of generations). Caloglossa monosticha Kamiya occurs only in Western Australia (WA) and is a P-type. Caloglossa ogasawaraensis Okamura occurs in WA, Northern Territory (NT), Queensland (QLD), New South Wales (NSW), Victoria (VIC) and South Australia (SA) and is for the most part a P-type in culture. A few isolates have bisexual gametophytes that are self-compatible, while most are unisexual. Caloglossa ogasawaraensis from Adelaide, SA and from Wilsons Promontory, VIC are new records for these States. In Australia, Caloglossa postiae (King et Puttock) Kamiya et King occurs in NSW, NT and QLD. All nine isolates are P-type. Isolates of Caloglossa leprieurii (Montagne) G. Martens from NSW, NT, QLD, Tasmania, VIC and New Zealand are P-type except for the freshwater isolates in which tetraspore germlings do not reproduce. In some isolates mixed-phase reproduction is seen with male gametophytes producing both viable spermatia and tetrasporangia and female gametophytes producing procarps and sori with non-dividing sporangia. All isolates of C. leprieurii irom Spencer Gulf and Gulf of St Vincent, SA and one isolate from QLD give rise to successive asexual generations of tetrasporophytes. Based on RuBisCO spacer DNA data an asexual life history has arisen several times in the C. leprieurii complex. The literature on apomixis, mixed-phase reproduction, bisexuality and sexual compatibility in red algae is surveyed.

Reproductive patterns of Caloglossa species (Delesseriaceae, Rhodophyta) from Australia and New Zealand: multiple origins of asexuality in C. leprieurii. Literature review on apomixis, mixed-phase, bisexuality and sexual compatibility

Reproductive patterns of Caloglossa species (Delesseriaceae, Rhodophyta) from Australia and New Zealand: multiple origins of asexuality in C. leprieurii. Literature review on apomixis, mixed-phase, bisexuality and sexual compatibility