Diversity Of Reproductive Modes Research Articles

Endocrine regulation of reproductive biology in echinoderms: An evolutionary perspective from closest marine invertebrate relatives to chordates

Echinoderms are a phylum of invertebrate deuterostomes, which contain echinoids, asteroids, holothuroids, crinoids, and ophiuroids. Echinoderms have special evolutionary position and unique characteristics, including pentamerous radial body structure, elaborate calcareous endoskeletons, and versatile water vascular system. Echinoderms exhibit extraordinarily diverse reproductive modes: asexual reproduction, sexual reproduction, sexual reversal, etc. Endocrine regulation plays important well-known roles in sex differentiation, gonadal development and maturation, gametogenesis, and reproductive behavior in vertebrates. However, the entire picture of reproductive endocrinology in echinoderms as an evolutionary model of the closest marine invertebrate relatives to chordates has not been revealed. Here, we reviewed previous and recent research progress on reproductive endocrinology in echinoderms, mainly including two sections: Sex steroids in echinoderms and neuropeptide regulation in echinoderm reproduction. This review introduces a variety of endocrine regulatory mechanisms in reproductive biology of echinoderms. It discusses the vertebrate-like sex steroids, putative steroidogenic pathway and metabolism, and reproduction-related neuropeptides. The review will provide a deeper understanding about endocrine regulatory mechanisms of gonadal development in lower deuterostomes and the application of endocrine control in economic echinoderm species in aquaculture.

Chromosome-level assembly, annotation and phylome of Pelobates cultripes, the western spadefoot toad.

Genomic resources for amphibians are still hugely under-represented in vertebrate genomic research, despite being a group of major interest for ecology, evolution and conservation. Amphibians constitute a highly threatened group of vertebrates, present a vast diversity in reproductive modes, are extremely diverse in morphology, occupy most ecoregions of the world, and present the widest range in genome sizes of any major group of vertebrates. We combined Illumina, Nanopore and Hi-C sequencing technologies to assemble a chromosome-level genome sequence for an anuran with a moderate genome size (assembly span 3.09 Gb); Pelobates cultripes, the western spadefoot toad. The genome has an N50 length of 330 Mb with 98.6% of the total sequence length assembled into 14 super scaffolds, and 87.7% complete BUSCO genes. We use published transcriptomic data to provide annotations, identifying 32,684 protein-coding genes. We also reconstruct the P. cultripes phylome and identify 2,527 gene expansions. We contribute the first draft of the genome of the western spadefoot toad, P. cultripes. This species represents a relatively basal lineage in the anuran tree with an interesting ecology and a high degree of developmental plasticity, and thus is an important resource for amphibian genomic research.

Natural history traits of the terrestrial breeding frog Batrachyla taeniata (Anura: Batrachylidae) in wet meadows of Patagonia

ABSTRACT Anurans have diverse reproductive modes, and some species of the austral forests of Patagonia have become specialized in terrestrial reproduction. Batrachyla taeniata is one of these species: it inhabits wetlands, lays its eggs during late summer and fall in terrestrial conditions, and has overwintering tadpoles. We studied a population of B. taeniata that breeds in wet meadows surrounded by native Patagonian forest and described its life cycle parameters (breeding phenology, egg development, and coexistence with other anurans) over two different years with contrasting hydroperiods. We also analyzed the effect of predatory insects on tadpole survival. Embryo development occurred in wet soil, which may be exposed to the sun or not, and took from 48 to 63 days to hatch. Free-swimming larvae overwintered in the wetland and coexisted in spring with pond-breeding anurans that varied in composition and abundance according to the wetland hydroperiod. The survivorship of B. taeniata tadpoles was significantly greater than that of P. thaul tadpoles when both were exposed to a free belostomatid predator. We consider that the terrestrial reproductive mode and life cycle of B. taeniata enable this species to survive in the variable biotic conditions determined by hydroperiod length.

Phylogenomic inference of species and subspecies diversity in the Palearctic salamander genus Salamandra

The salamander genus Salamandra is widespread across Europe, North Africa, and the Near East and is renowned for its conspicuous and polymorphic colouration and diversity of reproductive modes. The phylogenetic relationships within the genus, and especially in the highly polymorphic species S. salamandra, have been very challenging to elucidate, leaving its real evolutionary history and classification at species and subspecies levels a topic of debate and contention. However, the distribution of diversity and species delimitation within the genus are critically important for identifying evolutionarily significant units for conservation and management, especially in light of threats posed by the pathogenic chytrid fungus Batrachochytrium salamandrivorans that is causing massive declines of S. salamandra populations in central Europe. Here, we conducted a phylogenomic analysis from across the taxonomic and geographic breadth of the genus Salamandra in its entire range. Bayesian, maximum likelihood and network-based phylogenetic analyses of up to 4905 ddRADseq-loci (294,300 nucleotides of sequence) supported the distinctiveness of all currently recognised species (Salamandra algira, S. atra, S. corsica, S. infraimmaculata, S. lanzai, and S. salamandra), and all five species for which we have multiple exemplars were confirmed as monophyletic. Within S. salamandra, two main clades can be distinguished: one clade with the Apenninic subspecies S. s. gigliolii nested within the Iberian S. s. bernardezi/fastuosa; and a second clade comprising all other Iberian, Central and East European subspecies. Our analyses revealed that some of the currently recognized subspecies of S. salamandra are paraphyletic and may require taxonomic revision, with the Central- and Eastern-European subspecies all being poorly differentiated at the analysed genomic markers. Salamandra s. longirostris – sometimes considered a separate species – was nested within S. salamandra, consistent with its subspecies status. The relationships identified within and between Salamandra species provide valuable context for future systematic and biogeographic studies, and help elucidate critical evolutionary units for conservation and taxonomy.

Egg-laying site, fecundity and degree of sexual size dimorphism in frogs

AbstractFemale fecundity is an important selective force leading to female-biased sexual size dimorphism (SSD) in frogs. Because anurans exhibit diverse reproductive modes, we investigated whether variation in SSD and fecundity are related with oviposition site. We asked whether arboreal breeding species show pronounced female-biased SSD and if, paradoxically, females have lower fecundity because of the costs of carrying oocytes and amplectant males. Conversely, we tested whether species that deposit eggs in concealed sites show less pronounced SSD, because females do not carry males and space limitation may reduce female size and fecundity. Our results showed that, in general, males were approximately 20% smaller than females. However, for species with hidden oviposition sites, males and females exhibited more similar body sizes and arboreal hylids showed more pronounced female-biased SSD. Overall, fecundity was higher in aquatic breeders, as expected, but in hylids, fecundity was smaller in arboreal breeders, which suggests that arboreality may impose restrictions on fecundity. By analysing SSD in a broader and more specific lineage (Hylidae), we found that reproductive microhabitat may also influence female size and fecundity, playing an important role in the evolution of SSD in frogs at different evolutionary scales.

Climate and mating systems as drivers of global diversity of parental care in frogs

AbstractAimAmphibians exhibit unusually diverse reproductive modes, including a wide array of parental care strategies. The evolutionary drivers of this diversity, however, remain unclear. Here, we investigate three major factors that might predict interspecific variation in parental care strategies: climate, intrasexual selection and social environment. We hypothesize that some forms of care evolved to cope with harsh conditions, such as dry or unpredictable habitats. We contrast this prediction with the hypothesis that parental roles have co‐evolved with the social environment and mating systems.LocationGlobal.Major taxa studiedFrogs and toads (Amphibia: Anura).Time periodExtant taxa that represent c. 220 Myr of evolutionary history.MethodsUsing geographical and behavioural data for 971 species of frogs and toads that represent 45 anuran families, we quantified the global distribution of four forms of parenting separately for males and females: nest building, nest and/or tadpole attendance, carrying and nourishment. We used phylogenetic comparative analyses to investigate whether climate, social environment and mating systems predicted interspecific variation in parental care.ResultsOur results showed that climatic effects contribute to parental care diversity: in cool and humid climates the males provide offspring attendance, whereas in predictable temperatures endotrophy occurs, whereby the female provides all nutrients for the offspring until metamorphosis. In addition, we found other associations between mating systems and forms of parental care: uniparental clutch attendance by males is present in species with territorial defence, whereas cooperative nest building co‐occurs with sperm competition. The type of parental care is not associated with adult sex ratios.Main conclusionsNo specific form of care is associated with hostile environments; in fact, some forms of care occur in beneficial conditions, whereas others are used independently from the climate. Instead, parenting diversity has co‐evolved closely with mating systems in frogs.

Adaptive response of extreme epiphyte Tillandsia species (Bromeliaceae) is demonstrated by different sexual reproduction strategies in the Brazilian Chaco

AbstractSome epiphytes are adapted to extreme environments with the ability to survive drought as a result of their morphological (xeromorphism), anatomical (foliar trichomes or scales) and physiological features. In contrast to vegetative features, they may have diverse sexual reproductive strategies. Here we compared the flowering morphology, floral biology, breeding system and pollinators of Tillandsia duratii, T. loliacea and T. recurvifolia (Bromeliaceae) adapted to an extreme environment, the Brazilian Chaco. Tillandsia duratii and T. recurvifolia flower for 5–6 months, whereas T. loliaceae flowers for 11 months, mainly in the dry season, with low to high flowering overlap between them. Although these species generally show similar flowering morphology, they differ in size, colour, odour and/or floral functionality, suggesting non-sharing of pollinators among them. Bimodal pollination occurs in T. duratii (bees, moths other than hawkmoths) and T. recurvifolia (butterflies, hummingbirds); in T. loliacea, we recorded no floral visitors. Tillandsia recurvifolia is self-incompatible, has flowers which approach herkogamy and protandry, and depends on pollen vectors for fruit set. The other two species show reverse herkogamy, autonomous self-pollination and self-compatibility (mixed breeding system); therefore, pollinators are not required. The disparity in reproductive strategies among the three species suggests the possibility of selection for diverse modes of reproduction.

Asexual parasites and their extraordinary host ranges.

In diverse parasite taxa, from scale insects to root-knot nematodes, asexual lineages have exceptionally large host ranges, larger than those of their sexual relatives. Phylogenetic comparative studies of parasite taxa indicate that increases in host range and geographic range increase the probability of establishment of asexual lineages. At first pass, this convergence of traits appears counter-intuitive: intimate, antagonistic association with an enormous range of host taxa correlates with asexual reproduction, which should limit genetic variation within populations. Why would narrow host ranges favor sexual parasites and large host ranges favor asexual parasites? To take on this problem I link theory on ecological specialization to the two predominant hypotheses for the evolution of sex. I argue that both hypotheses predict a positive association between host range and the probability of invasion of asexual parasites, mediated either by variation in population size or in the strength of antagonistic coevolution. I also review hypotheses on colonization and the evolution of niche breadth in asexual lineages. I emphasize parasite taxa, with their diversity of reproductive modes and ecological strategies, as valuable assets in the hunt for solutions to the classic problems of the evolution of sex and geographic parthenogenesis.

Ovarian control and monitoring in amphibians

Amphibian evolution spans over 350 million years, consequently this taxonomic group displays a wide, complex array of physiological adaptations and their diverse modes of reproduction are a prime example. Reproduction can be affected by taxonomy, geographic and altitudinal distribution, and environmental factors. With some exceptions, amphibians can be categorized into discontinuous (strictly seasonal) and continuous breeders. Temperature and its close association with other proximate and genetic factors control reproduction via a tight relationship with circadian rhythms which drive genetic and hormonal responses to the environment. In recent times, the relationship of proximate factors and reproduction has directly or indirectly lead to the decline of this taxonomic group. Conservationists are tackling the rapid loss of species through a wide range of approaches including captive rescue. However, there is still much to be learned about the mechanisms of reproductive control and its requirements in order to fabricate species-appropriate captive environments that address a variety of reproductive strategies. As with other taxonomic groups, assisted reproductive technologies and other reproductive monitoring tools such as ultrasound, hormone analysis and body condition indices can assist conservationists in optimizing captive husbandry and breeding. In this review we discuss some of the mechanisms of ovarian control and the different tools being used to monitor female reproduction.

Reproductive Biology of the Critically Endangered Valcheta Frog,Pleurodema somuncurense(Anura: Leptodactylidae), from Patagonia, Argentina

Fil: Velasco, Melina Alicia. Consejo Nacional de Investigaciones Cientificas y Tecnicas; Argentina. Universidad Nacional de la Plata. Facultad de Ciencias Naturales y Museo. Division Zoologia de Vertebrados. Seccion Herpetologia; Argentina

The Insect Pathogens.

Fungi are the most common disease-causing agents of insects; aside from playing a crucial role in natural ecosystems, insect-killing fungi are being used as alternatives to chemical insecticides and as resources for biotechnology and pharmaceuticals. Some common experimentally tractable genera, such as Metarhizium spp., exemplify genetic diversity and dispersal because they contain numerous intraspecific variants with distinct environmental and insect host ranges. The availability of tools for molecular genetics and multiple sequenced genomes has made these fungi ideal experimental models for answering basic questions on the genetic and genomic processes behind adaptive phenotypes. For example, comparative genomics of entomopathogenic fungi has shown they exhibit diverse reproductive modes that often determine rates and patterns of genome evolution and are linked as cause or effect with pathogenic strategies. Fungal-insect pathogens represent lifestyle adaptations that evolved numerous times, and there are significant differences in host range and pathogenic strategies between the major groups. However, typically, spores landing on the cuticle produce appressoria and infection pegs that breach the cuticle using mechanical pressure and cuticle-degrading enzymes. Once inside the insect body cavity, fungal pathogens face a potent and comprehensively studied immune defense by which the host attempts to eliminate or reduce an infection. The Fungal Kingdom stands alone in the range, extent, and complexity of their manipulation of arthropod behavior. In part, this is because most only sporulate on cadavers, so they must ensure the dying host positions itself to allow efficient transmission.

A unique mating strategy without physical contact during fertilization in Bombay Night Frogs (Nyctibatrachus humayuni) with the description of a new form of amplexus and female call.

Anurans show the highest diversity in reproductive modes of all vertebrate taxa, with a variety of associated breeding behaviours. One striking feature of anuran reproduction is amplexus. During this process, in which the male clasps the female, both individuals’ cloacae are juxtaposed to ensure successful external fertilization. Several types of amplexus have evolved with the diversification of anurans, and secondary loss of amplexus has been reported in a few distantly related taxa. Within Nyctibatrachus, a genus endemic to the Western Ghats of India, normal axillary amplexus, a complete loss of amplexus, and intermediate forms of amplexus have all been suggested to occur, but many species remain unstudied. Here, we describe the reproductive behaviour of N. humayuni, including a new type of amplexus. The dorsal straddle, here defined as a loose form of contact in which the male sits on the dorsum of the female prior to oviposition but without clasping her, is previously unreported for anurans. When compared to known amplexus types, it most closely resembles the form of amplexus observed in Mantellinae. Furthermore, we prove that, opposed to the situation in most anurans, male semen release happens before egg deposition. We hypothesize that the male ejaculates on the female’s dorsum and that sperm subsequently runs from her back and hind legs before fertilizing the eggs. A second feature characterizing anuran breeding is the advertisement call, mostly produced solely by males. Despite recent descriptions of several new Nyctibatrachus species, few studies have explored their vocal repertoire. We describe both the male advertisement call and a female call for N. humayuni. The presence of a female call has not been reported within Nyctibatrachidae, and has been reported in less than 0.5% of anuran species. Altogether, our results highlight a striking diversity and several unique aspects of Nyctibatrachus breeding behaviour.

Cryobiology: principles, species conservation and benefits for coral reefs

Coral reefs are some of the oldest, most diverse and valuable ecosystems on Earth because they can support one-quarter of all marine life in our oceans. Despite their importance, the world’s coral reefs continue to be degraded at unprecedented rates by local and global threats that are warming and creating a more acidic ocean. This paper explores the reproductive challenges of coral for ex situ conservation, using IVF and cryopreservation, and our practical biobanking methods. Coral present challenges for cryopreservation because their reproductive period is often limited to a few nights yearly, they are mostly hermaphrodites with diverse modes of reproduction, including asexual reproduction (i.e. fragmentation and parthenogenesis) and sexual reproduction (i.e. self- and cross-fertilisation) and they express physiological toxins that can inhibit cryopreservation. We have banked spermatozoa from 12 coral species using the same field-hardy methods and have created new coral with thawed spermatozoa. In addition, we describe the cryopreservation of coral symbionts, whose physiology only permits the highest success seasonally. As part of a multidisciplinary conservation strategy, these collections may provide a major hedge against extinction for corals facing the damaging effects of climate change and loss of genetic diversity, and promise to help offset threats to our reefs worldwide.

The biology, ecology and conservation of elasmobranchs: recent advances and new frontiers.

The elasmobranchs (sharks, skates and rays) are cartilaginous fishes that are important predatory and scavenging species within aquatic ecosystems and as living resources to human societies globally in cultural, economic, health, biodiversity and conservation contexts. There are about 900 extant species, including the world’s largest fish, that arose sometime during c. 400 million years of independent evolution. Extant elasmobranchs possess an intriguing set of biological characteristics, including, for example, larger brains than other ectothermic vertebrates, slow growth and late age at maturity, internal fertilisation and female sperm storage, diverse reproductive modes including oviparity and viviparity, relatively low fecundity, and social systems with dominance hierarchies and segregation by sex and age. In some respects the behavioural, ecological and life-history strategies of elasmobranchs share certain characteristics that are arguably more similar to mammals and birds than other fishes. These attributes in addition to other more unique features have also made elasmobranchs important model organisms for developmental, biomedical and evolutionary research. Furthermore, elasmobranchs occupy a very broad range of habitats worldwide and have a key role in the functioning of ecosystems. Taken together this identifies elasmobranchs as an important group to study for advancing fundamental biological knowledge in key fields ranging from molecular genetics and genomics to physiology, and from behaviour to population and community ecology. However, life-history traits such as slow growth, late maturity and low fecundity make elasmobranchs less resilient to exploitation. The conservation of elasmobranch populations in the face of pressures such as the expansion of industrialised fisheries, the degradation or removal of habitats, and for many species little or no management, require novel solutions for sustainable exploitation. To approach these and other challenges a more detailed understanding of elasmobranch biology and ecology will be the firm foundation upon which new ideas and initiatives relevant to conservation are built. The aim of this symposium is to bring together scientists interested in the study of fundamental biological characteristics and the ecological and evolutionary relationships of elasmobranchs, including studies focusing on their population trends and conservation status. The topics covered will span levels of biological organisation from molecules and cells to systems biology, and from individuals, populations and up to communities and ecosystems. The meeting aims to highlight significant recent advances and will identify new and interdisciplinary areas likely to make a key impact in the near future for improved conservation.

Absence of consistent genetic differentiation among several morphs of Actinia (Actiniaria: Actiniidae) occurring in the Portuguese coast.

Actinia equina, the beadlet sea anemone, is a very labile species, displaying variable colour patterns, broad habitat choice and diverse modes of reproduction. Historically, studies using genetic markers such as allozymes and differences in habitat choice lead several authors to propose that different colour morphs could represent different species. One of the species defined was A. fragacea. In this paper, the relationships between brown, red and green colour morphs of A. equina and A. fragacea were studied, using two DNA fragments (one mitochondrial and one nuclear). Individuals were sampled from three different areas in Portugal separated by a maximum distance of 500 km. This is the first study applying direct sequencing of selected gene fragments to approach the validity of Actinia morphs as different genetic entities. The results show that, at least in the Portuguese coast, these colour morphs do not correspond to the two valid species recognized in the literature. The existence of cryptic species is discussed.

Rapid Identification of Both Sexes ofLeptocybe invasaFisher & La Salle (Hymenoptera: Eulophidae: Tetrastichinae): A Morphological Perspective

The blue gum chalcid, Leptocybe invasa Fisher & La Salle (Hymenoptera: Eulophidae: Tetrastichinae) is a global pest in Eucalyptus plantations. Diverse modes of reproduction (thelytokous and sexual reproduction) are one of the advantages for rapid colonization and outbreak of this species. Understanding the reproductive strategies is advantageous for us to explore the invasive mechanisms. Importantly, how to identify both sexes of the species in the field is a crucial issue before we explore their reproductive strategies. In the current study, indexes of rapid identification of both sexes in L. invasa were studied. Results showed that the obvious differences in both sexes were the antenna and body size. Male wasps have three anelli and four funicular segments, but in females there are four and three segments, respectively. The longer male antennae, which are covered with longer and finer hairs than in the female, was also observed. Body, expansus, head, and gaster sizes of female wasps were significantly larger than the males. These results are an important base not only for exploring the reproductive strategies, but also for controlling L. invasa.

Mud-packing frog: a novel breeding behaviour and parental care in a stream dwelling new species of Nyctibatrachus (Amphibia, Anura, Nyctibatrachidae).

Reproductive modes are diverse and unique in anurans. Selective pressures of evolution, ecology and environment are attributed to such diverse reproductive modes. Globally forty different reproductive modes in anurans have been described to date. The genus Nyctibatrachus has been recently revised and belongs to an ancient lineage of frog families in the Western Ghats of India. Species of this genus are known to exhibit mountain associated clade endemism and novel breeding behaviours. The purpose of this study is to present unique reproductive behaviour, oviposition and parental care in a new species Nyctibatrachus kumbara sp. nov. which is described in the paper. Nyctibatrachus kumbara sp. nov. is a medium sized stream dwelling frog. It is distinct from the congeners based on a suite of morphological characters and substantially divergent in DNA sequences of the mitochondrial 16S rRNA gene. Males exhibit parental care by mud packing the egg clutch. Such parental care has so far not been described from any other frog species worldwide. Besides this, we emphasize that three co-occurring congeneric species of Nyctibatrachus, namely N. jog, N. kempholeyensis and Nyctibatrachus kumbara sp. nov. from the study site differ in breeding behaviour, which could represent a case of reproductive character displacement. These three species are distinct in their size, call pattern, reproductive behaviour, maximum number of eggs in a clutch, oviposition and parental care, which was evident from the statistical analysis. The study throws light on the reproductive behaviour of Nyctibatrachus kumbara sp. nov. and associated species to understand the evolution and adaptation of reproductive modes of anurans in general, and Nyctibatrachus in particular from the Western Ghats.

Anurofauna da bacia do Rio Tijuco, Minas Gerais, Brasil e sua relação com taxocenoses de anfíbios do Cerrado e suas transições

A supressão de paisagens naturais tem dizimado populações e/ou reduzido a área de ocorrência de muitas espécies de anuros, caso do Cerrado, dado o grande potencial econômico e velocidade de descaracterização por atividades humanas. Por este motivo, inventários de espécies são extremamente necessários nessas regiões que dependendo do grau de alteração ambiental podem prejudicar as atividades de dispersão e reprodução dos anuros. Este estudo tem por objetivo determinar a riqueza e a diversidade de modos reprodutivos da anurofauna da bacia do Rio Tijuco e comparar a composição da taxocenose dessa região com taxocenoses de anfíbios de outras regiões do bioma Cerrado e da floresta estacional semidecidual anteriormente estudadas. O estudo foi desenvolvido ao longo da bacia do Rio Tijuco numa extensão aproximada de 80 km. Através de quatro métodos de amostragem, foram registradas 40 espécies de anuros pertencentes a seis famílias, o que corresponde a 20% das espécies de anuros registradas para o Cerrado. As condições da paisagem favorecem espécies mais tolerantes e adaptadas ao clima mais seco, o que reflete numa taxocenose que possui baixa proporção de modos reprodutivos e uma elevada representatividade de modos generalizados, dependentes de água ou com adaptações contra dessecação. Em comparação com outras 15 localidades verificou-se que a anurofauna do Cerrado é típica, com baixa similaridade taxonômica com ecossistemas adjacentes. A diversidade de habitats e a expressiva riqueza de espécies tornam a bacia do Rio Tijuco de grande importância para a conservação da anurofauna do bioma Cerrado.

Alternative reproductive modes of Atlantic forest frogs

Diversity in reproductive modes is well known in amphibians, mainly among anurans, which are characterized by a diversity in breeding biology that exceeds that of any other tetrapod. Currently, 39 reproductive modes are recognized among anurans and some species display more than one mode. The breeding biology of some Brazilian Atlantic forest anurans was investigated for this study. We observed unreported reproductive modes for six species, variability in the reproductive modes of individuals of the same species (whereby some individuals of a given population displayed unusual reproductive modes when the physical conditions of the breeding site were suboptimal), and variations within the modes. These observations suggest possible evolutionary steps for the reproductive modes. Anuran breeding biology seems to be more diverse than previously reported, and a character matrix could be constructed to describe the total range of variation of the anuran reproductive modes.

Diverse modes of reproduction in the marine free-living ciliate Glauconema trihymene

BackgroundMost free-living ciliates reproduce by equal fission or budding during vegetative growth. In certain ciliates, reproduction occurs inside the cyst wall, viz. reproductive cysts, but more complex reproductive strategies have generally been thought to be confined to parasitic or symbiotic species, e.g. Radiophrya spp.ResultsIn addition to equal fission, asymmetric binary division and reproductive cysts were discovered in the free-living bacterivorous scuticociliate Glauconema trihymene Thompson, 1966. Asymmetric division is an innate physiological state that can be induced by sufficient food, and the higher the food concentration, the longer the asymmetric division persists. During asymmetric division, nuclear and somatic structures divide with transiently arrested cytokinesis and variable positioning of macronuclei. Phylogenetic analysis, based on the small subunit of ribosomal DNA (SSU rDNA) sequences, showed that the G. trihymene isolate studied here nests with typical scuticociliates and is paraphyletic to both the symbiotic apostome and astome ciliates, some of which also produce progeny by asymmetric division.ConclusionsThe asymmetric division in G. trihymene has no precedent among undisturbed free-living ciliates. The coexistence of multiple modes of reproduction may represent a previously undescribed reproductive strategy for ciliates living on food patches in coastal waters. This may also be indicative of similar reproductive strategies among other polyphenic ciliates, which have not been intensively studied. Asymmetric division provides a special opportunity for studying ciliates' phenotypic plasticity and may also illuminate the origins of multicellularity.