Distribution,population structure and salinity tolerance of the invasive amphipod Gmelinoides fasciatus (Stebbing) in the Neva Estuary (Gulf of Finland,Baltic Sea)

In the early 1970s, the Baikalian amphipod Gmelinoides fasciatus (Stebbing) was intentionally introduced into several lakes in the Gulf of Finland basin in order to enhance fish production. By 1996, G. fasciatus successfully colonized the littoral zone of Lake Ladoga and, via the Neva River, invaded the Neva Bay, the freshwater part of the Neva Estuary. In 1999, G. fasciatus was first registered in the inner Neva Estuary, the very first record of the Baikalian amphipod in brackish waters of the Baltic Sea. Distribution, abundance, reproduction and population structure of G. fasciatus in the Neva Estuary were studied during 1998–2000. In some locations of the Neva Estuary, maximum densities of G. fasciatus reached 3500 ind. m⁻². In general, density and biomass of G. fasciatus in the freshwater part of the Neva Estuary were higher (around 1.5 fold) than in the brackish-water part. Fecundity of this amphipod averaged 10–20 eggs per female, depending on body size of females and season. In order to assess the possibility of further spread of G. fasciatus in the Baltic Sea, the salinity tolerance of this species was determined in a series of laboratory experiments. Our results showed that the invasive amphipod G. fasciatus is potentially able to colonize shallow coastal habitats of, for example, the Gulf of Bothnia, Gulf of Riga and other parts of the Baltic Sea with water salinities ranging from 1 to 5 psu.

     

Mating behavior of a Northern Italian population of <Emphasis Type="Italic">Fusarium verticillioides</Emphasis> associated with maize

Fusarium verticillioides, the most common causal organism of Fusarium stalk and ear rot of maize in Northern Italy, produces important mycotoxins such as fumonisins. Reproductive biology of F. verticillioides has been widely studied in numerous maize growing areas, but up to now no information is available on the mating behavior and genetic structure of this plant pathogen in Italy. Mating type and female fertility distribution and effective population number, N _e , were assessed for a population of 181 F. verticillioides strains isolated from three fields located in Lombardia region (Northern Italy) during 2007-2008 maize growing season. The ratio of MAT-1:MAT-2 was significantly different from the theoretical 1:1 ratio expected in an idealized population in which individuals mate at random. The frequency of hermaphroditic strains was 20 % of the total population. N _e for mating type was 89 % of the count (total population) and the N _e for male or hermaphrodite status was 55 %. The number of isolates that can function as the female parent limited N _e in the examined population. Under equilibrium cycle, assuming that female fertility has been lost due to selection and mutation rate during asexual reproduction, sexual reproduction needed to occur only once per 40 to 118 asexual generations to maintain this level of sexual fertility.     

Sex at the origin: an Asian population of the rice blast fungus Magnaporthe oryzae reproduces sexually

Sexual reproduction may be cryptic or facultative in fungi and therefore difficult to detect. Magnaporthe oryzae, which causes blast, the most damaging fungal disease of rice, is thought to originate from southeast Asia. It reproduces asexually in all rice‐growing regions. Sexual reproduction has been suspected in limited areas of southeast Asia, but has never been demonstrated in contemporary populations. We characterized several M. oryzae populations worldwide both biologically and genetically, to identify candidate populations for sexual reproduction. The sexual cycle of M. oryzae requires two strains of opposite mating types, at least one of which is female‐fertile, to come into contact. In one Chinese population, the two mating types were found to be present at similar frequencies and almost all strains were female‐fertile. Compatible strains from this population completed the sexual cycle in vitro and produced viable progenies. Genotypic richness and linkage disequilibrium data also supported the existence of sexual reproduction in this population. We resampled this population the following year, and the data obtained confirmed the presence of all the biological and genetic characteristics of sexual reproduction. In particular, a considerable genetic reshuffling of alleles was observed between the 2 years. Computer simulations confirmed that the observed genetic characteristics were unlikely to have arisen in the absence of recombination. We therefore concluded that a contemporary population of M. oryzae, pathogenic on rice, reproduces sexually in natura in southeast Asia. Our findings provide evidence for the loss of sexual reproduction by a fungal plant pathogen outside its centre of origin.     

Food composition and feeding rate of the lake Baikal invader <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis> (Stebbing, 1899) in Lake Ladoga

The feeding of the Baikal invader Gmelinoiudes fasciatus (Stebbing, 1899) in Lake Ladoga has been studied based on field and experimental data. The food spectrum of this amphipod was revealed; it includes many plant and animal species. Differences in the feeding of G. fasciatus living in different littoral biotopes were shown. The ecological and physiological rations of this species were determined. The effect that the bottom substrate has on the feeding rate of the amphipod was estimated. The introduction of G. fasciatus into Lake Ladoga provided a more adequate utilization of the energy accumulated by the primary producers and its transfer to higher trophic levels. This Baikal amphipod occupied a niche that had been, in effect, vacant. This made the invader capable of maintaining the quantitative parameters of its population at a very high level, and it largely reorganized the flow of matter and energy in the littoral zone of the lake.     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.     

Establishment of the Baikalian endemic amphipod Gmelinoides fasciatus Stebb. in Lake Ladoga

Gmelinoides fasciatus Stebb., a small amphipod from the Lake Baikal basin, was discovered in July 1988 in Lake Ladoga, the largest European lake. G. fasciatus likely invaded Lake Ladoga as a consequence of its intentional introduction, aimed at enhancing fish production, in some Karelian Isthmus lakes close to Lake Ladoga's western shore in the early 1970's. Benthos studies conducted in 1989 and 1990 revealed that G. fasciatus was well established in littoral communities along the western and northern shores of Lake Ladoga. G. fasciatus was the dominant species in these littoral communities and contributed over 70% of the macroinvertebrate biomass. The species was abundant in different macrophyte beds and stony littoral areas, both in heavily polluted and undisturbed sites. The maximum abundance and biomass of G. fasciatus was about 54,000 ind m^–2 and 160 g (wet wt.) m^–2. Negative impacts of G. fasciatus on native species have been observed. Studies are needed to identify the effects of G. fasciatus on the functioning of littoral communities and to predict and control the spread of this amphipod.     

Population structure and spatial distribution of <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis> (Stebb.) in the cooling reservoir of the Belovo Thermal Power Plant

The sex ratio and size-age structure of the Gmelinoides fasciatus (Stebb.) population in different areas of the cooling reservoir of the Belovo Thermal Power Plant (Kemerovo oblast) has been analyzed. The highest values of abundance and biomass of the species were recorded in areas with the lowest heating and lowest temperatures of the natural thermal regime. The invasion of G. fasciatus in the Belovo Reservoir has not caused the extinction of the indigenous species Gammarus lacustris Sars.     

Feeding selectivity,food assimilability and demand of the Baikal invader <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis> (Stebbing, 1899) in Lake Ladoga

Feeding selectivity and assimilability of the Baikal invader Gmelinoides fasciatus (Stebbing, 1899) in Lake Ladoga at feeding by various types of plant and animal food have been studied experimentally. The food demand has been assessed at different assimilation levels. Freshwater shrimp G. fasciatus has been shown to have distinct selective capacity with respect to different kinds of plant and animal food.     

Life cycle patterns of rotifers in Lake Peipsi

Life cycle strategies of rotifers in Lake Peipsi (Estonia) were examined. Bisexual (mictic) reproduction was detected in 26 species. Life cycle patterns were determined for 17 species. All three basic life cycle patterns were represented, with some evidence of intraspecific variability. The midcycle pattern, with mixis occurring near the population maximum, prevailed. Extended mixis was observed in littoral populations of Polyarthra luminosa and P. remata. Most species probably are monocyclic. Polyarthra dolichoptera and Synchaeta oblonga may be dicyclic. Mictic periods were not detected in Conochilus hippocrepis and C. unicornis, which probably are acyclic in L. Peipsi. Mictic ratios and egg ratios were calculated for some of the dominant species. The highest amictic egg ratios were observed slightly before or at the population maxima. High mictic ratios (1.0) indicated very intense bisexual reproduction in populations of A. fissa, P. dolichoptera and S. verrucosa. Definite periods of bisexual reproduction could not be distinguished in the rotifer community. During May–October, the spread of mictic reproduction merely reflects the general seasonal distribution of rotifers. Their life cycle strategies represent specific adaptations to unpredictability of their habitat. The results of the study confirm that mixis is an anticipatory event, and not a response to environment deterioration, or an ending of a rotifer population cycle.     

Life cycle of a small endemic goby Economidichthys pygmaeus (Family: Gobiidae) in Lake Pamvotis,northwestern Greece

The life history tactics of the small endemic freshwater goby Economidichhtys pygmaeus were examined in a Mediterranean shallow lake located in the northwestern part of Greece (Lake Pamvotis). The purpose of the study was to define the population structure, growth, length at sexual maturity, sex ratio, annual cycle of gonad development, spawning characteristics and fecundity. Fish samples were collected using a hand net from October 2006 to September 2007. The fish grew rapidly in summer after hatching, matured after their first winter, and bred from February to June. The length frequency distribution indicated the appearance of two cohorts per year. Juveniles occurred in samples from April to September. Maturity length (ML₅₀) of males and females were estimated to be 23.54 and 23.28 mm TL, respectively. Sexual dimorphism related to the male caudal fin was observed and males grew to greater lengths than females. The overall ratio of males to females was 1:1.35. Spawning was from February to June. Ripe females appeared to be multiple spawners, with oocytes at different stages of development. Size of oocytes changed during maturation and increased in diameter from about 0.125 to more than 0.9 mm. Absolute fecundity was positively related to TL and TW and varied between 59 and 309 eggs. After reproduction, the number of parental individuals decreased.     

Reproductive biology of the broad‐snout [Chondrostoma holmwoodii (Boulenger, 1896)] in Tahtalı Dam Lake, İzmir,Turkey

The study aimed to determine the reproductive characteristics of the broad‐snout Chondrostoma holmwoodii population (spawning season, number of eggs and first age of sexual maturity) in order to assess the population status in the Tahtal? Dam Lake reservoir. A total of 1208 specimens were examined of which 72.1% were females and 27.9% males (female:male ratio = 1 : 0.40); the percentage of males, however, declined in the higher age classes (IV and V). The monthly determined gonadosomatic index indicated that the peak reproduction period of the species is in April and early May. The study covered age classes I–V, with specimens at age II reaching maturity. The total number of eggs per female varied from 8000 to 41 700, the mean number of eggs increasing proportionally with age of the spawner. Egg diameters also increased during the 5‐month ripening period from December to April.     

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.     

Population structure and reproduction of <Emphasis Type="Italic">Eukrohnia bathypelagica</Emphasis> and <Emphasis Type="Italic">Eukrohnia bathyantarctica</Emphasis> in the Lazarev Sea,Southern Ocean

In the Lazarev Sea, Southern Ocean, I studied the population structure and reproduction of Eukrohnia bathypelagica and E. bathyantarctica during winter and summer. A lack of unimodality in their population structures indicated at least two generations building one population and a life cycle longer than 1 year. During both seasons all maturity stages were present; therefore, continuous reproduction is very probable. Extensive breeding seasons with several releases of eggs by one generation are assumed, as adults with empty marsupial sacs continue to build new ova. E. bathypelagica carried between 86 and 128 eggs in both marsupial sacs. E. bathyantarctica had only between 8 and 13 eggs. Although self-fertilization seemed at least to be possible in E. bathyantarctica, cross-fertilization appears to be more common in both species, as most individuals developed testes and ovaries consecutively.     

Sexual reproduction and life cycle of Ceratium furcoides (Dinophyceae) in situ in the lake Plußsee (F.R.)

The life cycle of Ceratium furcoides (Levander) Langhans which features sexual reproduction is described for the first time in situ from the lake Plußsee (northern West Germany). Anisogamic copulation occurs during the bloom period in July–August by incorporation of the male gamete by the female gamete, resulting in an overwintering planozygote. The planozygote develops into a benthic hypnozygote which germinates in spring to establish a planomeiocyte. The morphology and structure of the stages of the life cycle are described using light and scanning electron microscopy.     

AUXOSPORULATION IN CYCLOTELLA OCELLATA (BACILLARIOPHYCEAE) UNDER NATURAL AND EXPERIMENTAL CONDITIONS1

Growth and sexual reproduction in a population of Cyclotella ocellata Pantocseck were studied during one annual cycle in a reservoir and in short-term enclosure experiments performed in situ involving different nutrient conditions and concentrations of zooplankton species. Three phases of auxosporulation in this diatom were distinguishable morphologically: 1) preauxospore, from the beginning of zygote formation until the valves were longitudinally separated, 2) primary auxospore, when the zygote grew too large to fit inside the valves and before it reached its full size, and 3) mature auxospore, characterized by a well-developed, markedly scalloped edge. Under experimental and natural conditions, sexual reproduction was associated with changes in cell size. In the natural system, the auxospore appeared to act as a resting structure during conditions adverse for population growth. A threshold population of small cells appeared to be necessary for sexual reproduction in the natural system, whereas auxosporulation was associated with phosphorus fertilization in the enclosures. In both environments only cells smaller than 9.5 μm in diameter were capable of auxospore formation. Our results suggest that, once having reached the critical cell size, the factors that trigger sexual reproduction may depend on ambient environmental conditions.     

THE SEXUAL AND ASEXUAL LIFE CYCLE OF GLENODINIOPSIS STEINII (DINOPHYCEAE)

The sexual and asexual portions of the life cycle of Glenodiniopsis steinii were examined at both the light and scanning electron microscopic levels. Asexual reproduction by cell division occurs every 2 to 3 d under optimal conditions. Gamete formation and sexual reproduction can be induced by either nitrogen deprivation or aging of cultures. Fusion of gametes peaks about 10 hr into the light cycle and requires about 10 hr for completion. Cells remain motile during fusion but the zygote loses motility. The cell then undergoes changes that give it a thick-walled, warty appearance.     

The reproductive biology of <Emphasis Type="Italic">Nassarius festivus</Emphasis> (Powys, 1835) (Gastropoda: Nassariidae) in relation to seasonal changes in temperature and salinity in subtropical Hong Kong

The reproduction of Nassarius festivus (Gastropoda: Nassariidae) has been studied on three contrasting beaches in Hong Kong. There was no significant difference in shell heights between females and males of N. festivus at three study sites and the sex ratio was always female biased. The highest female sex ratio was recorded from Starfish Bay, followed by Tai Tam Bay and then Tai Mong Tsai. The sexual cycle of N. festivus is completed in 1 year in Hong Kong. It is a winter spawner with a reproductive period ranging from September to May. The gonads progressively mature in autumn with maturity being achieved in winter. Spring and summer are times of gonadal decline and gametogenesis initiation, respectively. The ranges in water temperature and salinity at which mature individuals occur were 13.7–28.0°C and 19.3–29.0‰, respectively. N. festivus produces between 16 and 46 eggs·capsule⁻¹. Over its short life span of 1.21 years, therefore, N. festivus attempts to maximize reproductive output. The eggs hatch as veligers which remain in the plankton for between 33 and 41 days. In terms of its reproductive biology, it is concluded that N. festivus behaves as a typical sub-tropical species with a mix of tropical and temperate characteristics seen in other nassariids from these latitudes.     

Geographic variability in the gonadal development and sexual ontogeny of Hemigymnus,Cheilinus and Oxycheilinus wrasses among Indo-Pacific coral reefs

Patterns of reproductive ontogeny in four species of coral reef wrasses (F: Labridae) Hemigymnus melapterus, Hemigymnus fasciatus, Cheilinus fasciatus and Oxycheilinus digramma were investigated. Populations of each species were sampled from two island groups of the central Great Barrier Reef (GBR), Australia, and from coral reefs in the central Philippines. These three sampling locations span 30° of latitude. The GBR and Philippine reefs experience biologically significant differences in water temperature, geography and human activity. The studied wrasses are effectively unfished in Australia but heavily fished in the Philippines. Gonad weights, histology and demographic data were obtained across the entire size and age range of H. melapterus, C. fasciatus and O. digramma from all locations. Analysis identified three processes of male recruitment: functional gonochorism and both forms of protogynous hermaphroditism, monandry and diandry. The expression of these distinct sexual ontogenies was locality dependent. Populations of H. melapterus, H. fasciatus, C. fasciatus and O. digramma on the GBR showed consistently uniform patterns of sexual ontogeny, with all species being exclusively monandric. H. melapterus, C. fasciatus and O. digramma in the Philippines displayed complex sexual ontogenies, with all species showing histological evidence of both diandry and functional gonochorism. Reproductive investment in gonadal tissue, and population sex structure, also differed between GBR and Philippine coral reefs. Philippine populations had substantially lower gonado-somatic indices than populations on the GBR. Nonetheless, Philippine populations matured more rapidly and displayed a protracted timing of sex change over a large size and age range. Thus, mature females appeared earlier and persisted later into ontogeny in the Philippines than on GBR reefs. Protracted timing of sex change on Philippine reefs is likely linked to the presence of primary males in the population, which is known to reduce the strength of selection for mature females to undergo sex change and become male. Hypotheses based on social structure of fish populations, environmental factors and evolutionary history were developed to account for the different patterns of sexual ontogeny in the focal wrasses.     

Demographic consequences of greater clonal than sexual reproduction in Dicentra canadensis

Clonality is a widespread life history trait in flowering plants that may be essential for population persistence, especially in environments where sexual reproduction is unpredictable. Frequent clonal reproduction, however, could hinder sexual reproduction by spatially aggregating ramets that compete with seedlings and reduce inter‐genet pollination. Nevertheless, the role of clonality in relation to variable sexual reproduction in population dynamics is often overlooked. We combined population matrix models and pollination experiments to compare the demographic contributions of clonal and sexual reproduction in three Dicentra canadensis populations, one in a well‐forested landscape and two in isolated forest remnants. We constructed stage‐based transition matrices from 3 years of census data to evaluate annual population growth rates, λ. We used loop analysis to evaluate the relative contribution of different reproductive pathways to λ. Despite strong temporal and spatial variation in seed set, populations generally showed stable growth rates. Although we detected some pollen limitation of seed set, manipulative pollination treatments did not affect population growth rates. Clonal reproduction contributed significantly more than sexual reproduction to population growth in the forest remnants. Only at the well‐forested site did sexual reproduction contribute as much as clonal reproduction to population growth. Flowering plants were more likely to transition to a smaller size class with reduced reproductive potential in the following year than similarly sized nonflowering plants, suggesting energy trade‐offs between sexual and clonal reproduction at the individual level. Seed production had negligible effects on growth and tuber production of individual plants. Our results demonstrate that clonal reproduction is vital for population persistence in a system where sexual reproduction is unpredictable. The bias toward clonality may be driven by low fitness returns for resource investment in sexual reproduction at the individual level. However, chronic failure in sexual reproduction may exacerbate the imbalance between sexual and clonal reproduction and eventually lead to irreversible loss of sex in the population.     

Differences in the effects of parental age on offspring life history between tropical and temperate populations of milkweed bugs (Oncopeltus spp.)

Summary We compare the effects of parental age on several offspring life history traits in two milkweed bug populations: a typically univoltine population ofOncopeltus fasciatus from the Sacramento Valley of California and a typically multivoltine population ofO. cingulifer from Monteverde, Costa Rica. Reared under identical conditions (27°C, 12 h light: 12 h dark photoperiod), these bugs exhibit significant differences in the effects of parental age on offspring life history. As they age,O. fasciatus females from Sacramento lay clutches of eggs of decreasing average weight and a decreasing proportion of their eggs produce offspring that survive to adulthood. Those offspring which do survive have a significantly faster developmental rate and females have a larger body size at adult eclosion. AsO. cingulifer from Monteverde age, they also produce lighter eggs, but there is no significant change in the offspring developmental rate, survival or female adult body size. We suggest that these results are largely explicable as the consequence of different parental investment strategies associated with the predictable relationship between parental age and time of season inO. fasciatus but not inO. cingulifer. AsO. fasciatus from Sacramento age, they may be increasing their investment per developing offspring so as to increase the probability that nymphs hatching late in the season will reach a prereproductive adult diapause before the first killing frost.