Appearance of males in a thelytokous strain of Milnesium cf. tardigradum (Tardigrada)

Tardigrades are generally gonochoristic. Many moss-dwelling species propagate by parthenogenesis, but heterogony has not yet been found. Milnesium tardigradum, a carnivorous tardigrade, also has both sexes, but males are usually rare and many populations appear to have only parthenogenetic reproduction. Since 2000, I have maintained a thelytokous strain of Milnesium cf. tardigradum that originated from one female. Individuals of this strain were thought to be all females, but here I report that males have emerged in this strain at a very low frequency. This is the first report of the appearance of males in parthenogenetic tardigrades. On the first pair of legs of some individuals, I observed the modified claws characteristic of males of this species. It is unknown whether these males can actually function in sexual reproduction; however, they might allow some possibility of genetic exchange among clonal populations. No environmental factors that generate males were determined.     

Going solo: discovery of the first parthenogenetic gordiid (Nematomorpha: Gordiida)

Despite the severe fitness costs associated with sexual reproduction, its persistence and pervasiveness among multicellular organisms testifies to its intrinsic, short-term advantages. However, the reproductive assurance hypothesis predicts selection favoring asexual reproduction in sparse populations and when mate finding is difficult. Difficulties in finding mates is especially common in parasites, whose life cycles involve multiple hosts, or being released from the host into the external environment where the parasite can find itself trapped without a sexual partner. To solve this problem and guarantee reproduction, parasites in numerous phyla have evolved reproductive strategies, as predicted by the reproductive assurance hypothesis, such as hermaphroditism or parthenogenesis. However, this type of strategy has not been reported from species in the phylum Nematomorpha, whose populations have often been described as sparse. A new Nematomorpha species, Paragordius obamai n. sp., was discovered from Kenya, Africa, and appears to have solved the problem of being trapped without a mate by eliminating the need for males. Paragordius obamai n. sp. represents the first and only known species within this phylum to reproduce asexually. To determine the mechanism of this mating strategy, we ruled out the involvement of reproduction manipulating endosymbionts by use of next generation sequencing data, thus suggesting that parthenogenesis is determined genetically and may have evolved as a means to assure reproduction. Since this new parthenogenetic species and a closely related gonochoristic North American congener, P. varius, are easy to propagate in the laboratory, these gordiids can be used as model systems to test hypotheses on the genetic advantages and disadvantages of asexual reproduction and the genetic determinants of reproductive strategies in parasites.     

Life history variation in Pisidium (Bivalvia: Pisidiidae)

Intraspecific variation in the maximum shell length of Pisidium is small in comparison with variation in the life span of the clams, from 4 mo to 4 yr. The use of shell length as a measure of size is complicated by large intraspecific variation in the weight-length relationship, a possible reflection of resource availability on the weight of the soft parts. Maximum embryo length is relatively constant (1 mm) in the majority of species but litter size is variable (1 to 40), generally increasing with parent size. The time of egg-laying is determined by the size of the clam, season, and previous episode of reproduction, while the timing of the release of embryos, depending also on embryonic growth rate, is additionally affected by temperature and oxygen conditions. In brief, the maximum adult and embryo lengths are relatively conservative traits while growth rates vary by an order of magnitude.
Reproductive effort, when measured by the ratio of litter weight to parent weight, may increase or decrease with parent length and age. Both semelparous and iteroparous populations are known in several species. We suggest that if maximum adult length can be reached by the first reproductive season (in a favourable environment) the population is semelparous, otherwise (in an unfavourable environment) the population is iteroparous. Gravid clams in semelparous populations continue to grow, and thus assimilation cannot be meaningfully partitioned into reproduction, somatic growth and maintenance.
Along a depth gradient from 8 to 65 m, maximum and mature shell lengths, embryo length and litter size varies significantly in Pisidium conventus subpopulations not distinguishable electrophoretically. Large clams in deep water produce large embryos but small litters, which is contrary to the general trend of increasing litter size with parent length. Low food availability that decreases juvenile growth rate may restrict litter size in this case.     

Molecular genetic evidence for parthenogenesis in the Burmese python,Python molurus bivittatus

Parthenogenesis among reptiles is rare. Only a few species have the ability to reproduce asexually. Most of these are obligate parthenogenetic species that consist (almost) entirely of females, which can reproduce solely through parthenogenesis. Rarer are sexual species that only sporadically reproduce through parthenogenesis. A female Python molurus bivittatus (Reptilia, Boidae) from the Artis Zoo, Amsterdam, produced eggs in five consecutive years that contained embryos while she was isolated from males. These eggs might be fertilized with stored sperm, or might be the product of parthenogenesis. Parthenogenesis has not been shown for the Boidae family before. We performed parentship analyses on the snake and seven of her embryos using microsatellites and AFLP. Four microsatellite loci developed for this species combined with three loci developed previously for different snake species revealed too little variation to discriminate between sperm retention and parthenogenesis. With AFLP we were able to confirm that the Artis Zoo female reproduced parthenogenetically. Because the offspring are genetically identical to their mother, whereas in previous studies on sporadic parthenogenesis in snakes a loss of genetic information was reported, we conclude that the meiotic pathways that produce the diploid egg cells are different.     

Parthenogenesis in an elasmobranch in the presence of conspecific males

Parthenogenesis has been observed in several elasmobranch species, primarily in public aquaria. The majority of cases of parthenogenesis have occurred either when females were held without males or once a male was removed from a female's habitat. Here we report a second instance of parthenogenesis in a zebra shark female that was housed with conspecific mature males. This study calls into question the conditions under which elasmobranch females undergo parthenogenesis.     

The meiofauna of artificial water-filled tree holes: colonization and bottom-up effects

The meiofaunal community of artificial water-filled tree holes was determined, and the bottom-up effects of different amounts of leaf litter on abundance and diversity were estimated. We assume a positive impact of leaf litter on meiofaunal abundances, species diversity, and trophic links. Plastic cups with different amounts of leaf litter were placed in a beech forest (Teutoburg Forest, Bielefeld, Germany) for 24 weeks. As early as 1 week later, the artificial tree holes were colonized by bdelloid rotifers, tardigrades, and nematodes. Rotifers were dominant throughout the experiment, followed by nematodes and tardigrades. The 29 nematode species that were identified included bacterial and hyphal feeders, with common species such as Plectus cirratus/accuminatus and Aphelenchoides parietinus predominating. Impacts of water volume (up to complete desiccation), pH, and O₂ on the meiofaunal community were not detected, whereas the addition of leaf litter resulted in bottom-up effects. Nematode abundance, especially that of bacterial feeders, and species number increased with increasing leaf input. The predatory nematode Prionchulus muscorum was found only in treatments containing high leaf content. Rotifer abundances were partly negatively affected by the amount of added leaves and, like tardigrades, showed a reversal in their correlation at higher leaf inputs. Our study revealed the fast colonization of small water bodies by meiofaunal organisms and the importance of passively distribution. Furthermore, the results provide a comparison with the meiofaunal community in lakes and soil.     

The Spermatozoon in Tardigrades: Evolution and Relationships with the Environment

The spermatozoon ultrastructure of seven species of tardigrades, one heterotardigrade echiniscid and six eutardigrades, was analysed and described. Tardigrade species were collected from freshwater sediments, moss and leaf litter. Aphylogenetic evaluation was made and relationships of spermatozoon shape and habitat were discussed. The spermatozoon morphology of the moss-dwelling echiniscid(heterotardigrade) was very similar to that found in marine species of the same order. This indicates that in Echiniscoididae change of habitat had little influence on sperm structural organisation. In contrast, eutardigrade spermatozoa have developed several different structural organisations that may be related to family and/or habitat.     

Current status of the tardigrada: evolution and ecology

The Tardigrada are bilaterally symmetrical micrometazoans withfour pairs of lobopod legs terminating in claws or sucking disks.They occupy a diversity of niches in marine, freshwater, andterrestrial environments throughout the world. Some have a cosmopolitandistribution, while others are endemic. About 900 species havebeen described thus far, but many more species are expectedas additional habitats are investigated. Most are less than1 mm in body length and are opaque or translucent, exhibitingcolors such as brown, green, orange, yellow, red, or pink inthe cuticle and/or gut. Marine species are more variable inbody shape and overall appearance and generally exhibit lowpopulation density with high species diversity. Reproductivemodes include sexual reproduction and parthenogenesis, but muchremains to be known about development. Tardigrades have a hemocoel-typeof fluid-filled body cavity, a complete digestive tract, anda lobed dorsal brain with a ventral nerve cord with fused ganglia.Recent molecular analyses and additional morphological studiesof the nervous system have confirmed the phylogenetic positionof tardigrades as a sister group of the arthropods. The abilityof tardigrades to undergo cryptobiosis has long intrigued scientists.Although tardigrades are active only when surrounded by a filmof water, they can enter latent states in response to desiccation(anhydrobiosis), temperature (cryobiosis), low oxygen (anoxybiosis),and salinity changes (osmobiosis). Cryptobiotic states aid indispersal.     

The Persistence of Facultative Parthenogenesis in Drosophila albomicans

Parthenogenesis has evolved independently in more than 10 Drosophila species. Most cases are tychoparthenogenesis, which is occasional or accidental parthenogenesis in normally bisexual species with a low hatching rate of eggs produced by virgin females; this form is presumed to be an early stage of parthenogenesis. To address how parthenogenesis and sexual reproduction coexist in Drosophila populations, we investigated several reproductive traits, including the fertility, parthenogenetic capability, diploidization mechanisms, and mating propensity of parthenogenetic D. albomicans. The fertility of mated parthenogenetic females was significantly higher than that of virgin females. The mated females could still produce parthenogenetic offspring but predominantly produced offspring by sexual reproduction. Both mated parthenogenetic females and their parthenogenetic-sexual descendants were capable of parthenogenesis. The alleles responsible for parthenogenesis can be propagated through both parthenogenesis and sexual reproduction. As diploidy is restored predominantly by gamete duplication, heterozygosity would be very low in parthenogenetic individuals. Hence, genetic variation in parthenogenetic genomes would result from sexual reproduction. The mating propensity of females after more than 20 years of isolation from males was decreased. If mutations reducing mating propensities could occur under male-limited conditions in natural populations, decreased mating propensity might accelerate tychoparthenogenesis through a positive feedback mechanism. This process provides an opportunity for the evolution of obligate parthenogenesis. Therefore, the persistence of facultative parthenogenesis may be an adaptive reproductive strategy in Drosophila when a few founders colonize a new niche or when small populations are distributed at the edge of a species'' range, consistent with models of geographical parthenogenesis.     

Parthenogenesis is self-destructive for scaled reptiles

Parthenogenesis is rare in nature. With 39 described true parthenogens, scaled reptiles (Squamata) are the only vertebrates that evolved this reproductive strategy. Parthenogenesis is ecologically advantageous in the short term, but the young age and rarity of parthenogenetic species indicate it is less advantageous in the long term. This suggests parthenogenesis is self-destructive: it arises often but is lost due to increased extinction rates, high rates of reversal or both. However, this role of parthenogenesis as a self-destructive trait remains unknown. We used a phylogeny of Squamata (5388 species), tree metrics, null simulations and macroevolutionary scenarios of trait diversification to address the factors that best explain the rarity of parthenogenetic species. We show that parthenogenesis can be considered as self-destructive, with high extinction rates mainly responsible for its rarity in nature. Since these parthenogenetic species occur, this trait should be ecologically relevant in the short term.     

The biology and ecology of lotic Tardigrada

• 1 Tardigrades comprise a micrometazoan phylum that is a sister group of the arthropods.
• 2 They are components of the meiobenthos in lotic habitats, and ≈ 50–70 species have been reported in such habitats world‐wide. Approximately 800 species have been identified from all marine, freshwater, and terrestrial habitats.
• 3 Taxonomy is based primarily on the morphology of the claws, buccal‐pharyngeal apparatus, cuticle and eggs.
• 4 Reproductive modes include sexual reproduction (amphimixis) and parthenogenesis. The sexual condition of individuals may be either gonochorism, unisexuality, or hermaphroditism. Moulting occurs throughout the life of the tardigrade.
• 5 Latent states (cryptobiosis, including encystment, anoxybiosis, cryobiosis, osmobiosis and anhydrobiosis) enable tardigrades to withstand unfavourable environmental conditions.
• 6 Population densities, life histories, dissemination and biogeography of freshwater species are poorly known.

     

Influence of spatial structure on the maintenance of sexual reproduction

We include spatial extension into a model for the maintenance of sexual reproduction introduced recently. The model is based on a broad spectrum of resources, which regrow slowly. Other key features of the model are that sexual reproduction sets in when resources become scarce and that only a few genotypes can coexist locally. The extension of the model to several patches in space is done in two different ways. Model A is based on central egg deposition and allows migration of juveniles into all patches. Model B has a one-dimensional array of patches with migration only between neighboring patches. The main findings are that wide dispersal favors asexuals, while for slower migration there is a wide range of parameter values for which sexually reproducing species always win against asexuals. These results are conform with major patterns for the distribution of parthenogenesis in animals and plants, i.e. the prevalence of parthenogenetic reproduction in minute species, which are easily dispersed by physical forces, such as protists and small metazoans including e.g. bdelloid rotifers, tardigrades and nematodes.     

Hermaphroditism in tardigrades

Evidence is presented to show the existence of hermaphroditism in tardigrades, a phenomenon hitherto unknown for this phylum: specimens were obtained containing male and female germ cells in maturation in the same gonad. Hermaphroditic animals have been found in a few species of Isohypsibius; in many species of other genera and also of Isohypsibius, there is gonochorism.     

The emerging phylogenetic pattern of parthenogenesis in snakes

Parthenogenesis occurs across a variety of vertebrate taxa. Within squamate reptiles (lizards and snakes), a group for which the largest number of cases has been documented, both obligate and facultative types of parthenogenesis exists, although the obligate form in snakes appears to be restricted to a single basal species of blind snake, Indotyphlops braminus. By contrast, a number of snake species that otherwise reproduce sexually have been found capable of facultative parthenogenesis. Because the original documentation of this phenomenon was restricted to subjects held in captivity and isolated from males, facultative parthenogenesis was attributed as a captive syndrome. However, its recent discovery in nature shifts the paradigm and identifies this form of reproduction as a potentially important feature of vertebrate evolution. In light of the growing number of documented cases of parthenogenesis, it is now possible to review the phylogenetic distribution in snakes and thus identify subtle variations and commonalities that may exist through the characterization of its emerging properties. Based on our findings, we propose partitioning facultative parthenogenesis in snakes into two categories, type A and type B, based on the sex of the progeny produced, their viability, sex chromosome morphology, and ploidy, as well as their phylogenetic position. Furthermore, we introduce a hypothesis (directionality of heterogamety hypothesis) to explain the production of female‐only parthenogens in basal alethinophidian snakes and male‐only parthenogens in caenophidian (advanced) snakes.     

Extensive variation in chromosome number and genome size in sexual and parthenogenetic species of the jumping‐bristletail genus Machilis (Archaeognatha)

Parthenogenesis in animals is often associated with polyploidy and restriction to extreme habitats or recently deglaciated areas. It has been hypothesized that benefits conferred by asexual reproduction and polyploidy are essential for colonizing these habitats. However, while evolutionary routes to parthenogenesis are manifold, study systems including polyploids are scarce in arthropods. The jumping‐bristletail genus Machilis (Insecta: Archaeognatha) includes both sexual and parthenogenetic species, and recently, the occurrence of polyploidy has been postulated. Here, we applied flow cytometry, karyotyping, and mitochondrial DNA sequencing to three sexual and five putatively parthenogenetic Eastern‐Alpine Machilis species to investigate whether (1) parthenogenesis originated once or multiply and (2) whether parthenogenesis is strictly associated with polyploidy. The mitochondrial phylogeny revealed that parthenogenesis evolved at least five times independently among Eastern‐Alpine representatives of this genus. One parthenogenetic species was exclusively triploid, while a second consisted of both diploid and triploid populations. The three other parthenogenetic species and all sexual species were diploid. Our results thus indicate that polyploidy can co‐occur with parthenogenesis, but that it was not mandatory for the emergence of parthenogenesis in Machilis. Overall, we found a weak negative correlation of monoploid genome size (Cx) and chromosome base number (x), and this connection is stronger among parthenogenetic species alone. Likewise, monoploid genome size decreased with elevation, and we therefore hypothesize that genome downsizing could have been crucial for the persistence of alpine Machilis species. Finally, we discuss the evolutionary consequences of intraspecific chromosomal rearrangements and the presence of B chromosomes. In doing so, we highlight the potential of Alpine Machilis species for research on chromosomal and genome‐size alterations during speciation.     

The Tardigrades of Emilia (Italy). III. Piane di Mocogno (Northern Apennines)

Samples of leaf litter, mosses and lichens were collected in a beech forest at Piane di Mocogno (Emilia, Italy), 1200 m asl. The tardigrade community of the leaf litter was similar to that found in this substrate in Italy and in the U.S.A. At Piane di Mocogno the species association within the leaf litter was different from that in mosses and lichens (which have similar fauna), confirming that very different communities characterise different substrates such as leaf litter and mosses. During this faunal analysis, six species of tardigrades were found for the first time in this area. The presence of Microhypsibius bertolanii (first record in leaf litter and in Italy) and Eohypsibius nadjae are note worthy. Ornamented eggs in an exuvium of Hypsibius cf.scabropygus were also recorded for the first time.     

Gonochorism vs. hermaphroditism: relationship between life history and fitness in three species of Ophryotrocha (Polychaeta: Dorvilleidae) with different forms of sexuality

1. The relationships between life history, fitness and sexuality, together with their ecological and evolutionary significance, has been analysed comparing the main life-history traits and demography in three closely related species belonging to the genus Ophryotrocha. The species are: the gonochoristic O. labronica, the simultaneous hermaphrodite O. diadema and the protandrous hermaphrodite O. puerilis. 2. Survivorship and reproductive data were collected weekly and were used to construct life tables and population projection matrices for each species and compare life-history characteristics. Elasticity, life-table response and decomposition analyses were performed to examine the relative contribution of fecundity and survivorship to differences in lamda between species. 3. The gonochoristic and hermaphroditic species differ in all the main life-history parameters and also in demographic characteristics. In particular the value of lamda, used commonly to express fitness, is markedly higher in the gonochoristic species while in terms of fitness simultaneous and sequential hermaphroditism are very similar. In the genus Ophryotrocha gonochorism currently represents the most widespread condition, being characteristic of the majority of the known species in the genus. 4. Given the demographic advantage ensured by gonochorism, it remains be understood why some species have retained simultaneous hermaphroditism and one has evolved a sequential type hermaphroditism; the most probable hypothesis is correlated with the density of the species in natural habitats.     

First molecular genetic evidence for automictic parthenogenesis in cockroaches

Parthenogenesis is an asexual mode of reproduction that plays an important role in the evolution of sex, sociality, and reproduction strategies in insects. Some species of cockroach exhibit thelytoky, a type of parthenogenesis in which female offspring are produced without fertilization. However, the cytological and genetic mecha? nisms of parthenogenesis in cockroaches are not well understood. Here we provide the first molecular genetic evidence that cockroaches can reproduce through automixis. Using the American cockroach Periplaneta aniericana, we performed microsatellite analysis to investigate the genetic relationship between parthenogenetically produced nymphs and the parent virgin females, and found that all parthenogenetic offspring were homozygous for autosomal microsatellite markers, whereas the female parents were heterozygous. In addition, flow cytometry analysis revealed that the parthenogenetic offspring were diploid. Taken together, our results demonstrate that P. americana exhibits automixis-type thelytoky, in which diploidy is restored by gamete duplication or terminal fusion. These findings highlight the unique reproduction strategies of cockroaches, which are more varied than was previously recognized.     

Parthenogenesis in Mites and Ticks (Arachnida: Acari)

SYNOPSIS. Parthenogenesis is discussed according to type (arrhenotoky,thelytoky, deuteroLoky, artificial parthenogenesis, gynogenesis)and presence in various acarine taxa. Evidence for parthenogenesisis based on rearing and/or cytological studies. Arrhenotokyoccurs in Mesostigmata, Prostigmata and Astigmata and usuallydoes not occur sporadically throughout these taxa; when presentit operates in many closely related species as the major typeof reproduction in certain genera, subfamilies or families.Thelytoky is present in Mesostigmata, Metastigmata, Prostigmata,Astigmata and Cryptostigmata, but in no case is the major modeof reproduction in higher taxa, i.e. subfamilies, families.Its sporadic occurrence may or may not be accompanied by polyploidy.Deuterotoky occurs in one acarine group (Listrophoridae) andartificial parthenogenesis is possible in ticks (Metastigmata).Gynogenesis is present in several species of haplo-diploid Mesostigmata(some Dermanyssidae, Phytoseiidae), although it is usually consideredin connection with thelytoky. Current studies of the effectsof ploidy level on morphological variation indicate greatercharacter variability in diploid females than in haploid males.In some species of ticks thelytoky is expressed differentiallyeven on an intraspecific level. One geographic strain of a speciesmay have the ability to reproduce thelytokously, but if matedproduces males and females.     

The Inheritance of apomixis in Poa pratensis confirms a five locus model with differences in gene expressivity and penetrance

The genetic control of apomixis was studied in numerous segregating progenies originated from intercrossing and selfing of obligate sexual and facultative apomictic parents in Poa pratensis by means of the flow cytometric seed screen. The data support a novel model with five major genes required to control asexual seed formation: the Apospory initiator (Ait) gene, the Apospory preventer (Apv) gene, a Megaspore development (Mdv) gene, the Parthenogenesis initiator (Pit) gene, and the Parthenogenesis preventer (Ppv) gene. Differences in expressivity and interactions of these genes are responsible for the wide variation of the mode of reproduction. Apospory and parthenogenesis as well as the initiator and preventer genes of these components segregate independently. The genotypes with the highest expressivity of apospory and parthenogenesis were assigned as Ait-/apvapv/Pit-/ppvppv, those with intermediate expressivity as Ait-/Apv-/Pit-/Ppv-, and those with low expressivity as aitait/apvapv/pitpit/ppvppv. Among the self progenies of obligate sexual individuals, plants with a low capacity for apospory and/or parthenogenesis occurred, indicating that the sexual parents were heterozygous for the preventer genes and homozygous for the recessive initiator alleles (aitait/Apv-/pitpit/Ppv-). The dominant allele Ait exhibits incomplete penetrance. The degree of expressivity of apospory and parthenogenesis was constant among several harvest years of F1 plants.