The life-cycles of four amphipod species in the Kattegat

Populations of four species of Gammaridae (Crustacea, Amphipoda) were sampled over 15 months in the south-western Kattegat at Sangstrup Klint (Denmark). Chaetogammarus marinus, C. stoerensis and Gammarus oceanicus had annual life-cycles, while G. salinus most likely had two generations per year. Mature C. marinus females were found from March to July, mature C. stoerensis females from March to June, mature G. oceanicus females from December to early June and mature G. salinus females from late November to late July. The theoretical maximum number of clutches were estimated at five for C. marinus , five plus for C. stoerensis , and six for G. oceanicus and both winter and summer reproducing G. salinus . Spring was the period of main recruitment in all populations, with additional recruitment during summer in G. salinus .
The geographical patterns of the reproductive strategies are discussed in relation to environmental factors and interspecific competition.     

An experimental study on the effects of polymorphiasis in common eider ducklings.

Eight common eider (Somateria mollissima) ducklings were experimentally infected from 1 June through 13 June, 1995 with acanthocephalans (Polymorphus minutus) by allowing the birds to feed on Gammarus spp. (Gammarus oceanicus, G. salinus, G. zaddachi, and G. lacustris) containing acanthocephalan cystacanths. Uninfected Gammarus spp. were fed to a control group of seven ducklings. No mortality of ducklings occurred during the experiment. However, the infected ducklings gained weight more slowly than the control birds. After the 2 wk study period, the mean serum concentrations of total protein, albumin, beta-globulin, gamma-globulin, fructosamine and creatine kinase were lower in the infected group than in the controls. The mean (+/-SE) number of acanthocephalans in the intestine of the infected ducklings was 21 (+/-4). The parasites were attached to the mucosa of the posterior small intestine of the infected ducklings with a mixed inflammatory reaction consisting of heterophils and mononuclear lymphocytes surrounding the attachment sites.     

The relationship between range size and niche breadth: a test using five species of Gammarus (Amphipoda)

Data for five closely related species of gammarid crustaceans are used to examine interspecific relationships between the breadth of fundamental tolerance or capacity and geographical range size. Gammarus duebeni is, almost without exception, the most tolerant species and that with the best physiological performance. Although there is some limited variation, the remaining species can be ranked broadly in the sequence G. zaddachi  > G. salinus  >  G. oceanicus > G. locusta . The wide tolerance and high performance of G. duebeni is associated with the occupation of a wider range of environmental 'types' than any other of the species. In terms of geographical range size, the species can be ranked from most to least widespread in the sequence G. oceanicus  > G. duebeni  >  G. zaddachi  >  G. salinus  >  G. locusta . This provides little support for Brown's hypothesis, or the argument that the more widely distributed species within a taxonomic assemblage also tend to have the widest fundamental niches. However, if marine ( G. oceanicus and G. locusta ) and estuarine ( G. duebeni , G. zaddachi , G. salinus) species are considered separately, then in each case the species with the largest geographical range is also the most tolerant/best performer. In this sense, the jack-of-all-trades is the master-of-all, rather than the master-of-none.     

Zur Ökophysiologie,Sexualität und Populationsgenetik litoraler Gammaridea — ein Überblick

Comparative investigations on the physiological capacities in the euryhaline amphipodsGammarus locusta, G. oceanicus, G. salinus, G. zaddachi andG. duebeni were reviewed. In order to assess the adaptations of these species to the abiotic conditions of their environment, the following criteria were examined: oxygen consumption in relation to ambient salinity and temperature levels, respiratory responses following osmotic stress, resistance capacities to oxygen deficiency, resistance to aerial exposure and the simultaneous presence of hydrogen sulphide. Covering the range from marine to typically brackish-water inhabitants, the 5 species show adaptive responses in the above-mentioned order. Respiration is less intensely modified by external factors, and oxygen consumption decreases. Accompanied by faster rates of acclimation to new steady states of performance, resistance capacities increase. The significance of the findings obtained is discussed in relation to the environmental requirements of the amphipods considered. Based on breeding experiments, the sex-determining systems reported thus far inGammarus species are outlined. As demonstrated inG. duebeni, a more or less pronounced influence of external factors such as photoperiod may become effective. A preponderance of males was noted when offspring were raised under long-day photoperiods, whereas females prevailed under short-day conditions. In terms of the critical daylength, the light per day was estimated as being between 13 and 14 h (Elbe estuary population). Feminizing microporidians (Octosporea effeminans, Thelohania herediteria), which are transovarially transmitted, can interfere with the system of sex determination and sex differentiation of the host. As reflected in variousG. duebeni populations, they cause a maternally transferred sex-ratio condition by the production of all-female broods, thereby mimicking extrachromosomal inheritance. An increase of the salinity level to 25–30‰ results in a disappearance ofO. effeminans. In both microsporidians, long exposure to low temperatures (≤4°C) produces eggs which are not all parasitized. Furthermore, intersexuality can be induced by changing environmental factors. Microsporidian species have no influence on sex differentiation inG. duebeni celticus, G. salinus, G. locusta andG. pulex. Patterns of relative electrophoretic mobilities of proteins and the distribution of allele frequencies at polymorphic gene loci can be utilized for species diagnosis and for the evaluation of the relationships between different taxa, particularly at and below the species level. As exemplified by studies on several gammarids from marine, brackish and freshwater environments, inter- and infraspecific gene-enzyme variation is described. Electrophoretic investigations on natural populations of the euryhaline amphipodsG. zaddachi, G. salinus, G. tigrinus and others from different geographic areas provided evidence of considerable biochemical genetic variation. InTalitrus saltator- andTalorchestia deshayesii-populations the extent of variability based on micro-and macrogeographic aspects is illustrated. The large-scale genetic divergence is demonstrated by comparison of samples obtained from the Baltic, North, Atlantic and northern Mediterranean Seas.      

Phylogenetic Relationships of Five Species of the Amphipod Genus Gammarus

The phylogenetic relationships of five amphipod species of the genus Gummarus: G. oceanicus, G. locusta, G. salinus, G. zaddachi and G. duebeni are presented based on starch gel electrophoresis of 12 enzymes determined by 17 loci. The five species separate into three groups: G. locusta and G. duebeni each represent a distinct group while G. oceanicus, G. salinus and G. zaddachi together form a third group. This grouping results when the number of diagnostic loci, as well as when genetic distance between species pairs, are considered. The results compare well with morphological and, to some extent, ecological data.     

Electrophoretic approach to the biochemical systematics of gammarids

By utilizing the techniques for electrophoretic separation of proteins by vertical starch gels, the biochemical systematics of 10 Gammaridae species obtained from marine, brackish and freshwater habitats was studied. They includedChaetogammarus marinus, Gammarus zaddachi, G. salinus, G. oceanicus, G. tigrinus, G. chevreuxi, G. locusta, G. duebeni duebeni, G. d. celticus, G. pulex pulex, andG. fossarum. For comparison of electrophoretic mobilities selected enzymes (phosphoglucose isomerase, glutamate oxalacetate transaminase, arginine phosphokinase, hexokinase, leucine amino peptidase, mannose 6-phosphate isomerase) were assayed. They were used as diagnostic characters in terms of electrophoretic identities or diversities of most frequent alleles at polymorphic gene loci. These criteria could be applied to estimate intrageneric enzymic variation and degrees of genetic relatedness between the crustacean amphipod species under consideration, thereby complementing traditional morphological classification.     

Reproductive biology of Gambusia affinis holbrooki Baird and Girard, Xiphophorus helleri (Gunther) and X. maculatus (Heckel) (Pisces; Poeciliidae) in Queensland, Australia

The reproductive biology of three introduced Poeciliidae, Gambusia affinis holbrooki Baird and Girard, Xiphophorus helleri (Günther) and X. maculatus (Heckel) was studied over a 14-month period in Brisbane, Queensland. Photoperiod and water temperature determined the length of the breeding season in G. affinis , which produced up to nine broods between August and March, with the peak of reproductive activity in October. Fecundity was linearly related to body weight in G. affinis (mean fecundity = 22.78 ± 1.02, range = 3–108 offspring per brood).
Xiphophorus helleri bred from August to May, producing eight to nine broods, with peak reproductive activity from October to December. Fecundity was curvilinearly related to body weight (mean fecundity = 60.15 ± 3.8, range = 7–183 offspring per brood. Xiphophorus maculatus bred between September and March, producing five to six broods during the study period. Fecundity was linearly related to standard length (s.l.) (mean fecundity = 27.27 ± 2.4, range = 3–92 offspring per brood). Although the reproductive strategy of the three species differs, all have the capacity to build up large populations in south-eastern Queensland. Their present distributions and relative abundance are discussed.     

Congeneric amphipods show differing abilities to maintain metabolic rates with latitude

Metabolic variability across latitudinal populations of gammarid amphipods was examined in the summer by determining whole-animal rates of oxygen uptake (M(o)?) in four species with overlapping distribution patterns in the northeast Atlantic and Arctic oceans. Comparisons were made between an arctic/boreal species, Gammarus setosus, a subarctic/boreal species, Gammarus oceanicus, a boreal/temperate species, Gammarus duebeni duebeni, and a temperate species, Gammarus locusta. Measurements included acclimatized M(o)? in all four species and M(o)? after acclimation to 10°C in two populations of G. oceanicus and G. locusta. In G. oceanicus, acclimatized M(o)? declined with latitude (13° to 5°C) so that metabolic rates were lower in subarctic (79°N) relative to temperate (58°N) populations and similar to the values in G. setosus at 79°N. Consequently, there was no evidence for metabolic rate compensation in the colder-water, high-latitude populations in the summer. Further examination of the specific effects of temperature revealed similarities in M(o)? between populations of G. oceanicus acclimated at 10°C and similarities in thermal sensitivity (Q(10)) and activation energies (E(a)) on exposure to acute temperature change. In sharp contrast, there was no variation in summer acclimatized M(o)? with latitude in either G. d. duebeni between 48° and 70°N or G. locusta between 38° and 53°N. Instead, the two species maintained relatively high metabolic rates across latitudes, which were associated in G. locusta with differences in M(o)? and with Q(10) and E(a) values in amphipods acclimated at 10°C. The ability to compensate metabolic rate with latitude in the summer suggests greater metabolic flexibility, which predicts a greater capacity for survival during climate change of the temperate/boreal over the subarctic and arctic gammarid species.     

Selection of nest sites by a hole-nesting duck, the Goldeneye Bucephala clangula

Nest site preferences were examined for a population of Goldeneye Ducks breeding in nest boxes in Värmland, central Sweden. Some nest boxes were occupied more often than others even if females returning to the same nest box were excluded from the analysis. Nest boxes located higher up trees were occupied more often than those close to the ground and some spatial 'cluster groups' of boxes were occupied more often than others. Otherwise nest site prefernces were not related to any measured physical attributes of the boxes. Prefernces for nest boxes seemed to be based mainly on a tendency for females to select those that had been occupied by other females in the preceding year, especially if they had bred successfully. As a result of this, the occupancy of nest boxes was not random over years but rather progressed in a series of runs; a period of consecutive years in which a box was occupied was followed by a period of years in which it was empty.
There were reproductive consequences for these prefernces in that females occupying preferred boxes were less likely to lose their clutch to a predator. These females also bred earlier in the year and produced larger clutches and broods than females breeding in other boxes.     

Adaptive variation in sex determination in a crustacean, Gammarus duebeni

In some animals sex is determined after conception by environmental factors (environmental sex determination. ESD). In the amphipod Gammarus duebeni sex is reportedly determined by photoperiod: there is a higher proportion of males in broods reared under long-day than under short-day photoperiods. It has been proposed that this is an adaptive response to seasonal population dynamics. A test of the hypothesis would be to demonstrate changes in the degree to which sex is determined by the environment in populations from different latitudes with different dynamics. This study reports such a test. Environmental response is significantly less strong in a southern population with a long breeding season than in a northern one in which breeding is seasonally restricted. Moreover, the threshold of the ESD cue for male or female determination is not defined when the breeding season is weakly seasonal. There is a broad correlation between latitude (and hence breeding seasonality) and the strength of ESD response across a series of population studies. Similarities between the Gammarus system of sex determination and that of the Atlantic silverside Menidia menidia, a fish with thermal ESD, are discussed.     

Osmotic and ionic haemolymph concentrations in the Baltic Sea amphipod Gammarus oceanicus in relation to water salinity

The aim of this work was to determine the osmotic and ionic (Na(+), K(+), Ca(2+), Mg(2+) and Cl(-)) haemolymph concentrations in Gammarus oceanicus at different salinity levels. Being a species of marine origin it inhabits brackish waters of the Baltic Sea. G. oceanicus specimens were collected in January 2003 from the Gulf of Gdansk (salinity 7 psu). The animals were gradually acclimated to eight different salinity levels (5, 7, 14, 20, 25, 30, 35 and 41 psu) at a temperature of 5 degrees C and 100% oxygen saturation. The haemolymph osmolalities correlated positively with external salinity, from 545.4+/-17.3 mOsm in 5 psu to 1185.9+/-34.6 mOsm in 41 psu. G. oceanicus hyperregulated within the 5-31.5 psu range; above 31.5 psu it hyporegulated its body fluids in comparison to the external medium. At 31.5 psu (1017 mOsm) the haemolymph concentration of G. oceanicus was isoosmotic with the habitat. The haemolymph concentrations of all the studied ions, except K(+), correlated positively with their concentrations at the various salinity.     

Annual cycle in female Three-spined sticklebacks (Gasterosteus aculeatus L.) from an upland and lowland population

The annual cycles in female Three-spined sticklebacks from an upland (275 m) and a lowland (25 m) population were compared. The average length and weight of the lowland females were greater, but in both populations growth was checked in winter and again in the breeding season. Both populations also showed a winter decline in condition followed by a sharp increase in spring associated with the maturation of the ovaries. The hepato-somatic index of the females also increased during the spring, but the dry matter content of the liver declined during the spring and early summer. Lowland females had larger livers than the upland fish. In both populations average gonadosomatic index increased steadily throughout the winter, then abruptly in spring. The lowland population bred from May to early August, but the upland population bred only in May.
The energy content of the carcases of females was lowest in winter and again during the breeding season whereas the ash content showed maxima at these two periods. Lipid and glycogen analyses of the carcase, liver and ovaries also suggested that winter and the breeding season were periods of depletion from the carcase and liver, but not from the ovaries.
The observations indicate that the ovaries are to some extent insulated from fluctuations in resource availability. This supports experimental studies which indicate that egg production in the female stickleback can be subsidised by depletion from the carcase and liver when necessary.     

Two species of feminizing microsporidian parasite coexist in populations of Gammarus duebeni

The amphipod crustacean Gammarus duebeni hosts two species of vertically transmitted microsporidian parasites, Nosema granulosis and Microsporidium sp. A. Here it is demonstrated that these co-occurring parasite species both cause infected females to produce female-biased broods. A survey of European G. duebeni populations demonstrates that these two parasites co-occur in six of 10 populations. These findings contrast with the theoretical prediction that two vertically transmitted feminizing parasites should not coexist in a panmictic population of susceptible hosts at equilibrium. Possible explanations for the co-occurrence of the two feminizing microsporidia in G. duebeni include the recent invasion of a new parasite, horizontal transmission of one or both parasites and the spread of alleles for resistance to the dominant parasite in host populations.     

Increased mammalian predators and climate ch1ange predict declines in breeding success and density of Capercaillie <Emphasis Type="Italic">tetrao urogallus,</Emphasis> an old stand specialist,in fragmented Scottish forests

Low breeding success has been associated with declines in population and range of the re-introduced Capercaillie Tetrao urogallus in Scotland. Annual breeding success from 26 Scottish forests surveyed between 1991 and 2009 averaged only 0.6 chicks per female, the lowest rate recorded in 16 previous studies. Reduced breeding success was due to proportionally fewer females rearing chicks rather than a reduction in brood size. Birds bred less well in Perthshire at the southern edge of the range, where declines in indices of female and male density were highest. Only at the core of the range (Strathspey), where birds bred better, were female densities stable. Two weather variables, April temperature in the pre-breeding period (APRTEMP), and temperature at chick hatch in June (HATCHTEMP), increased over the study period. Indices of Pine Marten Martes martes increased 3.9-fold since 1995, and those of Red Fox Vulpes vulpes by 2.2-fold, whereas those of Carrion Crow Corvus corone, raptors and forest floor vegetation showed no change. Neither forest type nor forest ground vegetation appeared to influence breeding success. Instead, females reared more chicks in years when hatch time in June was drier, and in forests with lower marten and crow indices. In addition, more females reared broods in years when Aprils were cooler. Brood size was unaffected by any of the measured variables. Densities of adult birds were lower in forests with higher fox indices. Increased predation of clutches and chicks by martens and crows within these small, fragmented forests, as well as changes in climate, may explain reductions in breeding success and hence contribute to continued declines. Successful conservation of Capercaillie in remaining Strathspey strongholds may require better predator management, including a licensed removal of martens to test the hypotheses that martens contribute to reduced breeding success. This short-medium term approach will inform longer-term predator management policies and complement aspirations to increase the area and connectivity of forest habitat to benefit Capercaillie in-part through mitigating against any impact of increasing forest-edge predators.     

Shelter site use and secondary sex ratios in the woodlice Armadillidium vulgare and Porcellio scaber (Crustacea: Isopoda)

In grassland populations of the woodlice Armadillidium vulgare and Porcellio scaber (Latreille) the sex ratio in hand collections from beneath wooden offcuts and roofing tiles varied through the breeding season. The proportion of males was higher in these collections from surface refugia than in independent estimates of the population sex ratio taken from soil cores. Collections became female-biased in June and July when most females were carrying broods, most notably in P. scaber . It is suggested that males use surface shelter between foraging events. A. vulgare males are particularly abundant under boards in May as they become active on the soil surface in order to locate mates. Females may find the higher and more equable temperatures in refugia attractive because they reduce brood development time but the cost of this advantage is the risk of failure to balance moisture requirements. These patterns of habitat use by the sexes have important implications for several population processes, the evolution of life-history tactics and the development of mating systems in these species.     

Reproductive cycle and growth of Phyllopteryx taeniolatus

In this study, 36 males, 33 females and 15 juveniles of the common or weedy seadragon Phyllopteryx taeniolatus , a syngnathid fish endemic to the waters of southern Australia, were identified using visual implant fluorescent elastomer tags and pattern of appendages and their reproductive cycle and growth was studied from May 2001 to June 2002. Brooding males were present for a period of >6 months, from June–July to early‐January. The percentage of males that were pregnant in the studied areas peaked in November–December, near the end of the breeding season. Two pregnancies were reported for some males. Ten recruits were identified from November to June. Growth rates for young‐of‐the‐year, young of the previous year, males and females were measured. The von Bertalanffy growth constant K was estimated at 0·098 month⁻¹.     

All-female broods in Hypolimnas bolina (L.) A re-survey of West Fiji after 60 years

In the Xymphalid butterfly Hypolimnas bolina (L.) H. W. Simmonds in 1921 demonstrated, by breeding experiments, the occurrence on the islands of West Fiji of all-female broods, and he compared their frequency with that of normal bisexual broods in the same localities. In two areas he found only unisexual families (14 of them in Suva and one on Vanua Levuj, in one island (Ovalau) only a single bisexual brood and in the other two islands, Taveuni and Kandavu, one bisexual and one of each type respectively. Nearly 60 years, and probably at least 150 generations later, we found that all-female broods were still present in four of the five islands re-investigated, and breeding showed that in these four (which included Suva) the proportion of the two types of female was approximately equal. In the fifth island only a bisexual family was bred. The only marked change between 1921 and 1980 was in the Suva area, where Simmonds found far more unisexual females than we did.
By carrying out spermatophore counts in many of our wild females we found that almost all of them had been mated, the majority of them on one occasion only. Though the data are scanty, they suggest that there is no wastage of females and that the population in 1980 was stable for the two types of brood.
Reasons for our findings are discussed, particularly in the light of the Heuch model, but we conclude that no entirely satisfactory explanation has yet been given for the persistence of all-female broods. Again, there is so far no explanation of the mechanism though we favour a cytoplasmic factor.
We feel that the status of H, bolina as a mimic of species of Euploea is less assured as the result of our survey.     

Long-term population increase of black-browed albatrosses <Emphasis Type="Italic">Thalassarche melanophrys</Emphasis> at Heard Island, 1947/1948 – 2000/2001

The breeding population of the black-browed albatross Thalassarche melanophrys has increased at Heard Island since the first census data were obtained in 1947/1948. Four breeding localities are known, and all populations have increased in the period 1947/1948-2000/2001. The breeding population is estimated to have been approximately 200 pairs in 1947/1948. Based on 2000/2001 census data, the population has increased to a minimum of approximately 600 pairs over the 53 years. Two mechanisms, that of increased prey availability through scavenging discards from trawlers operating within their foraging range, and climatic amelioration, are proposed as hypotheses for this increase.     

Age of first breeding, dispersal and survival of Red Kites Milvus milvus in Wales

Twenty-nine Red Kites, that were tagged as nestlings in Wales, bred for the first time at 2–7 years old (mean 3–6). A minimum of 41% of tagged young had entered the breeding population by the 7th year. Individuals moved up to 22 km between birthplace and breeding place and, having bred, most stayed in the same locality from year to year. No differences in these various respects were found between the sexes, though two long movements by territorial birds both involved females. Annual survival of adult breeders was estimated at around 95%.     

Spring phenology and timing of breeding in short‐distance migrant birds: phenotypic responses and offspring recruitment patterns in common goldeneyes

Understanding how organisms adjust breeding dates to exploit resources that affect fitness can provide insights into impacts of climate change on avian demography. For instance, mismatches have been reported in long‐distance migrant bird species when environmental cues experienced during spring migration are decoupled from conditions on breeding grounds. Short‐distance migrant bird species that store reproductive nutrients prior to breeding may avoid or buffer adverse phenological effects. Furthermore, reduced short‐term reproductive success could be offset by higher future recruitment of surviving offspring. We evaluated whether recruitment of locally‐hatched female offspring was related to hatching date alone or strength of mismatched breeding date for 405 individually‐marked adult female common goldeneyes Bucephala clangula (a short‐distance migrant) and their ducklings from a site in central Finland where ice‐out date has advanced by ～ 2 weeks over 24 yr. Path analyses revealed that older, early‐nesting females with good body condition and larger broods recruited the most female offspring. Offspring recruitment decreased strongly among females that bred late relative to other females in the population each year; the extent of mismatched breeding date, i.e. hatching date scaled to annual ice‐out date, was less influential. Overall, most females advanced breeding dates when ice‐out occurred earlier in spring, but some females exhibited greater flexibility in response to ice‐out conditions than did others. In general, directional selection favoured early breeding over a wide range of ice‐out dates. Our results seem most consistent with a hypothesis that some short‐distance migrant species like goldeneyes have the capacity to track and respond appropriately to changing environmental conditions prior to onset of breeding.