Genetic structure in a progenetic trematode: signs of cryptic species with contrasting reproductive strategies

Complexes of cryptic species are rapidly being discovered in many parasite taxa, including trematodes. However, after they are found, cryptic species are rarely distinguished from each other with respect to key ecological or life history traits. In this study, we applied an integrative taxonomic approach to the discovery of cryptic species within Stegodexamene anguillae, a facultatively progenetic trematode common throughout New Zealand. The presence of cryptic species was determined by the genetic divergence found in the mitochondrial cytochrome c oxidase I gene, the 16S rRNA gene and the nuclear 28S gene, warranting recognition of two distinct species and indicating a possible third species. Speciation was not associated with geographic distribution or microhabitat within the second intermediate host; however frequency of the progenetic reproductive strategy (and the truncated life cycle associated with it) was significantly greater in one of the lineages. Therefore, two lines of evidence, molecular and ecological, support the distinction between these two species and suggest scenarios for their divergence.     

Alternative reproductive strategies in the progenetic trematode Coitocaecum parvum: comparison of selfing and mating worms

The progenetic opecoelid trematode Coitocaecum parvum can reproduce either precociously by selfing in its second intermediate amphipod host or by mating in its normal definitive fish host. In this study, we describe and compare the infection parameters and some life history traits of both egg-producing worms and non-egg producing worms in both their second intermediate and definitive hosts. We showed that 58% of worms start to produce eggs while still in the amphipod. The relative abundance of progenetic worms increased with amphipod size, and egg-producing worms achieved greater size in amphipods than in fish. These 2 findings support the reproductive insurance hypothesis. No difference in size was revealed between eggs produced in the amphipods and those produced in the fish. Although more information is needed to thoroughly assess the respective costs and benefits of selfing and mating in this species, our conclusion is that adopting progenesis may have few, if any, long-term negative consequences for the parasite.     

Geographic variation in thermal sensitivity of early life traits in a widespread reptile

Taxa with large geographic distributions generally encompass diverse macroclimatic conditions, potentially requiring local adaptation and/or phenotypic plasticity to match their phenotypes to differing environments. These eco‐evolutionary processes are of particular interest in organisms with traits that are directly affected by temperature, such as embryonic development in oviparous ectotherms. Here we examine the spatial distribution of fitness‐related early life phenotypes across the range of a widespread vertebrate, the painted turtle (Chrysemys picta). We quantified embryonic and hatchling traits from seven locations (in Idaho, Minnesota, Oregon, Illinois, Nebraska, Kansas, and New Mexico) after incubating eggs under constant conditions across a series of environmentally relevant temperatures. Thermal reaction norms for incubation duration and hatchling mass varied among locations under this common‐garden experiment, indicating genetic differentiation or pre‐ovulatory maternal effects. However, latitude, a commonly used proxy for geographic variation, was not a strong predictor of these geographic differences. Our findings suggest that this macroclimatic proxy may be an unreliable surrogate for microclimatic conditions experienced locally in nests. Instead, complex interactions between abiotic and biotic factors likely drive among‐population phenotypic variation in this system. Understanding spatial variation in key life‐history traits provides an important perspective on adaptation to contemporary and future climatic conditions.     

Geographic body size variation in the periodical cicadas Magicicada: implications for life cycle divergence and local adaptation

Seven species in three species groups (Decim, Cassini and Decula) of periodical cicadas (Magicicada) occupy a wide latitudinal range in the eastern United States. To clarify how adult body size, a key trait affecting fitness, varies geographically with climate conditions and life cycle, we analysed the relationships of population mean head width to geographic variables (latitude, longitude, altitude), habitat annual mean temperature (AMT), life cycle and species differences. Within species, body size was larger in females than males and decreased with increasing latitude (and decreasing habitat AMT), following the converse Bergmann's rule. For the pair of recently diverged 13‐ and 17‐year species in each group, 13‐year cicadas were equal in size or slightly smaller on average than their 17‐year counterparts despite their shorter developmental time. This fact suggests that, under the same climatic conditions, 17‐year cicadas have lowered growth rates compared to their 13‐years counterparts, allowing 13‐year cicadas with faster growth rates to achieve body sizes equivalent to those of their 17‐year counterparts at the same locations. However, in the Decim group, which includes two 13‐year species, the more southerly, anciently diverged 13‐year species (Magicicada tredecim) was characterized by a larger body size than the other, more northerly 13‐ and 17‐year species, suggesting that local adaptation in warmer habitats may ultimately lead to evolution of larger body sizes. Our results demonstrate how geographic clines in body size may be maintained in sister species possessing different life cycles.     

A brachylaimid trematode from marsupial mice with observations on its life cycle

A species of digenetic trematode, Brachylaime antechini (Family Brachylaimidae), is described from adult worms recovered from the intestine and rectum of two species of marsupial mice, Antechinus stuartii(10 of 56 infected) and Antechinus swainsonii (2 of 5 infected). On experimental evidence unencysted metacercariae from the kidney of a terrestrial gastropod, Strangesta capillacea, are considered to be larval stages of Brachylaime antechini. Miracidia, sporocysts and cercariae have not been recovered, but circumstantial evidence suggests that Strangesta capillacea fulfils the role of first intermediate host. Brachylaime antechini most closely resembles Brachylaime chiapensis, a parasite of deer mice in Mexico.     

The trematode Lecthodolfusia anatina (Pleurogenidae) and its life cycle in Primor'e

The trematiode Lecthodolfusia anatina Kchotenovsky, 1967 is redescribed. Experimental study has shown, that in the Primorye land, the first intermedial host of this trematode is a mollusk Boreoelona contortrix ussuriensis, and the second ones are larvae of the may fly Ecdyonurus aurarius and caddis flies Semblis phaenoides and Groerodes sp. Adult worm were reared in chickens.     

The morphology,life-cycle and systematic position of Orthetrotrema monostomum Macy & Basch, 1972, a progenetic trematode

The life-cycle of Orthetrotrema monostomum, a progenetic trematode occurring in the haemocoel of dragonfly naiads is described. The snail Thiara tuberculata acts as the intermediate host. The cercaria is xiphidiocercous, monostomate and belongs to the Microcotylous group. The course of development of the cercaria into a mature adult fluke is followed in the haemocoel of dragon-fly naiads which had been exposed to cercariae. On the basis of intraspecific variations noted in the adult flukes obtained experimentally from dragon-fly naiads, O. longicaeca Hanumantha Rao & Swamy, 1975 is synonymised with O. monostomum. Evidence is provided for the inclusion of the genus Orthetrotrema in the family Eumegacetidae. The diagnosis of the family Eumegacetidae is amended.     

Individual variation in movement throughout the life cycle of a stream-dwelling salmonid fish

Stream-dwelling fish populations have long served as important models of animal movement. Populations of adult stream-dwelling fishes are generally composed of a mix of relatively sedentary and mobile individuals. However, we do not know whether this pattern that we typically observe among adults is indicative of patterns of movement that occur throughout the life cycle. Therefore, we do not know whether we can apply these patterns to understanding or predicting processes such as migration and thus the potential for the evolution of genetic differences among populations. We test the general hypothesis that patterns of movement throughout the life cycle are consistent with patterns of movement inferred by indirect genetic methods and, more specifically, that the characteristics of the mobile fraction of the population are consistent with patterns of genetic differentiation. We used parentage analyses to infer the movements of alevin brook charr (Salvelinus fontinalis) in Freshwater River, Newfoundland, Canada, and a capture-recapture study of one cohort in this population to infer movement throughout the rest of the life cycle. We found that alevins move large distances shortly after emergence, primarily in the downstream direction, and that the population is composed of a mix of relatively sedentary and mobile individuals throughout all other intervals of the life cycle. In contrast, when we considered movements of individuals first captured as juveniles and eventually recovered as reproductively mature adults, we found relatively large and uniform distributions of net movement distance. Thus, heterogeneity in individual movement of adults is not representative of patterns of movement throughout the life cycle and therefore may provide only limited inference of population-level processes such as gene flow.     

Genotypic and phenotypic variation in transmission traits of a complex life cycle parasite

Characterizing genetic variation in parasite transmission traits and its contribution to parasite vigor is essential for understanding the evolution of parasite life‐history traits. We measured genetic variation in output, activity, survival, and infection success of clonal transmission stages (cercaria larvae) of a complex life cycle parasite (Diplostomum pseudospathaceum). We further tested if variation in host nutritional stage had an effect on these traits by keeping hosts on limited or ad libitum diet. The traits we measured were highly variable among parasite genotypes indicating significant genetic variation in these life‐history traits. Traits were also phenotypically variable, for example, there was significant variation in the measured traits over time within each genotype. However, host nutritional stage had no effect on the parasite traits suggesting that a short‐term reduction in host resources was not limiting the cercarial output or performance. Overall, these results suggest significant interclonal and phenotypic variation in parasite transmission traits that are not affected by host nutritional status.     

Lack of seasonal variation in the life-history strategies of the trematode Coitocaecum parvum: no apparent environmental effect

Parasites with complex life cycles have developed numerous and very diverse adaptations to increase the likelihood of completing this cycle. For example, some parasites can abbreviate their life cycles by skipping the definitive host and reproducing inside their intermediate host. The resulting shorter life cycle is clearly advantageous when definitive hosts are absent or rare. In species where life-cycle abbreviation is facultative, this strategy should be adopted in response to seasonally variable environmental conditions. The hermaphroditic trematode Coitocaecum parvum is able to mature precociously (progenesis), and produce eggs by selfing while still inside its amphipod second intermediate host. Several environmental factors such as fish definitive host density and water temperature are known to influence the life-history strategy adopted by laboratory raised C. parvum. Here we document the seasonal variation of environmental parameters and its association with the proportion of progenetic individuals in a parasite population in its natural environment. We found obvious seasonal patterns in both water temperature and C. parvum host densities. However, despite being temporally variable, the proportion of progenetic C. parvum individuals was not correlated with any single parameter. The results show that C. parvum life-history strategy is not as flexible as previously thought. It is possible that the parasite's natural environment contains so many layers of heterogeneity that C. parvum does not possess the ability to adjust its life-history strategy to accurately match the current conditions.     

Geographic variability of the life cycle in Pterostichus melanarius (Coleoptera, Carabidae)

In Pterostichus melanarius, the seasonal dynamics of activity is characterized by two peaks (in mid-June-early July and in late July-mid-August). In southwestern Moldova, these peaks were registered twenty days later than in Moscow Province. In both regions, larvae and adults hibernate. Both hibernated immature adults and adults that had already bred before wintering start reproduction simultaneously. In hibernated immature individuals, the average number of eggs per female constituted 1.5 times that in the postgenerative females. In the steppe zone, the total number of eggs laid by immature females was closed to that laid by previously bred females, while in the southern taiga zone, hibernated immature females laid a greater number of eggs. In the steppe zone, females of a new generation lay a somewhat lower number of eggs in comparison with hibernated immature females, whereas the total number of laid eggs is significantly higher. In the southern taiga zone, both the average number of laid eggs and their total number are equal in females of a new generation and hibernated immature females. In Moldova, the majority of individuals of a new generation breed in the current season, whereas in Moscow Prov., about half the individuals hibernate without breeding. Over the largest part of the range, the life cycle of P. melanarius combines the one-year development with hibernating larvae with two-year development with the hibernating immature and postgenerative adults. The two subpopulation groups, wintering at different phases of the ontogenesis, are totally isolated at the northern border of the range. Within each subpopulation., individuals are characterized by the biennial life cycle; as a result, the polyvariant character of the life cycle turns into the monovariant one. We suggest that such a life cycle should be designated as the compensatory monovariant cycle, and populations, in which it is realized, as byside populations (from the English “side by side”).     

The life form concept and the use in the analysis of life cycle evolutionary strategies

The life form is a generalized morphoecological characteristic of an animal giving an idea of the organism as a whole, its position and function and functional role in the ecosystem. This characteristic is inherent to a species or to a group of congeneric species (for applied goal it is better to use a genus, not species) considered in the framework of higher taxon (from family to type). The principal contradiction of life form concept is determined by the existence of ontogenetic stages and changes of life forms during the whole life of individual. It is usually assumed that the concept of life form should be applied only to the adult stage, thus ignoring the integral character of the life cycle as indivisible unit of selection, evolution and functioning in ecosystem. We propose that a morphologically specific ontogenetic of a given species should be used as an elementary lowest unit in the classification of life forms. Thus it can be considered as integrated internally structured morphoecological unit in time and multidimensional space of abiotic and biotic environmental factors. As an example we describe the types of reproductive strategies and classification of elementary (ontogenetic) life forms in cephalopods. We present characteristics of the life cycle of some typical cephalopod species inhabiting different biotopes and having different models of locomotion, feeding, reproduction and development.     

Egg development, nymphal growth and life cycle strategies in Plecoptera

Interspecific and intraspecific differences in egg development, nymphal growth and the life cycles of Fennoscandian stoneflies are examined in relation to zoogeographical aspects and evolutionary trends. Data on other European and North American species are also considered.
Egg development shows considerable diversity, the main categories being ovovivipary, egg development with diapause and non-diapause development. Egg development in species in the non-diapause category can be classified as eurythermal, cold stenothermal or warm stenothermal. Temperature has a marked effect on nymphal growth of some species but not others. The distribution of several species can be explained by their temperature relationships, either in the egg or nymphal stage. Several species modify their life cycle to fit local climatic conditions. In some species, such as Leuctra hippopus , environmental selection has been so strong that they have probably recently evolved into subspecies and/or species in northern Europe.
Morphological and ecological variations in the Plecoptera have facilitated their colonization of the variety of freshwater biotopes that occur in Fennoscandia.     

Alternative life cycle strategies of Megalodiscus temperatus in tadpoles and metamorphosed anurans

Megalodiscus temperatus (Stafford, 1905) is a common paramphistome trematode of North American amphibians with a two host life cycle and has been reported to infect frogs and rarely tadpoles. In this study we document the alternative life cycle strategy of M. temperatus in tadpoles and metamorphosed anurans. We show through field work and experimental infections that M. temperatus can establish in both anuran life stages and worms become gravid and release eggs in both tadpoles and metamorphosed frogs. However, worms exhibit differences in route of infection, development, egg production, and diet in tadpoles and metamorphosed anurans. These alternative life history strategies of M. temperatus suggest different selective pressures on the development and reproductive success of these worms in tadpoles and metamorphosed anurans, and we discuss the evolutionary avenues for and constraints on amphibian trematode life cycles presented by these two different anuran life stages.     

Geographic variation in Giardia karyotypes

Chromosomes of 41 stocks of Giardia duodenalis derived from humans and 14 stocks from other animal species were analysed by field inversion gel electrophoresis (FIGE). These stocks have two predominant karyotypes as judged by FIGE which appear to fit a geographic distribution. Under FIGE conditions used to optimize the detection of size variation in Giardia chromosomes, five or six major chromosomes could be identified. Most of the stocks derived from North America have three major chromosomes smaller than 800 kb while most of the Australian stocks have four. A few exceptions, and minor variations, of these karyotypes were observed. It was estimated that not all of the DNA entered the gel, the remainder being trapped conformations or very large chromosomes. Karyotypes of Giardia stocks from different animal hosts and human sources within a geographical region are similar.