Population dynamics in a Danish metapopulation of spadefoot toads Pelobates fuscus

If an amphibian species has a high degree of site fidelity to its natal pond, subpopulations do not mix, and this may have implications: the subpopulations may get different because of genetic drift, and their status may be affected due to stochastic events. During a four-year period (1994–1997), demographic parameters of a Danish metapopulation consisting of ca 1000 adult Pelobates fuscus in five subpopulations were investigated with the aim of quantifying exchange of individuals in subpopulations and potential differences in demographic parameters, and to clarify population regulating mechanisms. This was done through the use of drift fences surrounding the five breeding ponds, individual marking (PIT-tagging) and weight recordings of the adult females. Metamorphs were recorded when leaving the pond. There was no significant differences between ponds in sex ratio, eggs laid per female, fraction of females breeding and survival of eggs to froglets and of adults. However, there was a significant difference in juvenile survival (52% in one pond and 31% in the others). Sex ratio was 1.70 males:1 female. Mean number of eggs laid per female in the pond was 1762. The number of eggs laid was age-specific, fecundity peaking at age five. Larval survival rate was density dependent: decreasing in an overcompensating fashion at high densities. Exchange rate between ponds was low; 1.09% of the marked and recaptured adult population was found to change pond during the study period. The low exchange rate was confirmed in a simple translocation experiment. This population shows features of a source-sink metapopulation where the identity of the source pond changes due to succession. It is presumably regulated either in the larval stage by density dependent mechanisms (in the subpopulations with many adults) or in the adult stage by density independent mechanisms (in the subpopulations with few adults).     

Can the limited marsupium space be a limiting factor for Syngnathus abaster females? Insights from a population with size-assortative mating

1. Some syngnathid species show varying degrees of sex role reversal aside from male pregnancy, with females competing for access to mates and sometimes presenting conspicuous secondary sexual characters. Among other variables, brooding space constraints are usually considered a key element in female reproductive success, contributing strongly to the observed morphological and behavioural sexual differences. Nevertheless, a close relationship between sex role reversal and male brooding space limitation has not yet been accurately demonstrated in field studies. 2. The present work, conducted over two consecutive breeding seasons in a wild population of the sex role-reversed pipefish Syngnathus abaster, simultaneously analysed egg number and occupied space, as well as the free area in the male's marsupium. The number of eggs that would fit in the observed unoccupied space was estimated. 3. Contrary to what would be expected, given the marked sexual dimorphism observed in the population studied, where females were larger and more colourful, male brooding space did not appear to limit female reproduction as neither large nor small individuals presented a fully occupied pouch. Interestingly, the largest unoccupied areas of marsupium were found in the larger individuals, although they received more and larger eggs. Laboratory data also showed that larger females lay larger eggs. 4. Together, these results suggest the existence of assortative mating, which may result from: (i) the reluctance of larger males (which tend not to receive small eggs usually laid by small females) to mate with lower quality females, even at the expense of a smaller number of offspring; or (ii) female-female competition, which might strongly reduce the hypothesis of a small female mating with a large male. The potential impact of temperature on reproduction and population dynamics is also discussed in the light of ongoing climatic changes.     

Male breeding behaviour and mate acquisition in the Common toad, Bufo bufo

The breeding behaviour of a Common toad ( Bufo bufo ) population was studied at a breeding pond on Portland, Dorset. Major toad movements towards the breeding pond occurred on warm, wet nights when the temperature did not fall below 6°C. Males arrived earlier than females resulting in an initially exaggerated sex ratio which then slowly reduced as the breeding season progressed. No size assortative pairing was found although large males were more successful at mating than small males. Although males demonstrated no ESS (evolutionary stable strategy) in searching for a female between sites within the pond evidence suggesting the existence of one between the pond and an undefined area surrounding it was found.     

Acquisition of Salmonella flora by turtle hatchlings on commercial turtle farms

A commercial turtle pond in South Louisiana was studied to identify the mechanism by which turtle hatchlings acquire Salmonella flora. The visceral organs and mature eggs removed from 31 adult gravid female turtles over the course of two egg-laying seasons and from 37 adult females during one winter dormant period were examined bacteriologically for Salmonella. Pond water, egg nest soil, and hatchlings produced by eggs removed from the oviducts and nest soil were also tested. Eighty-eight turtles hatched from eggs removed from the oviducts of 15 turtles at necropsy did not excrete or harbor systemically Salmonella, nor were these pathogens isolated from ovarian tissue or immature eggs. The findings suggest transovarian transmission of these pathogens does not occur frequently. Turtles hatched from eggs retrieved from soil nests 1 to 2 h after deposition harbor and excrete these organisms. This result coupled with the isolation of these pathogens from the cloaca, colon contents, and bursal fluid from 18 females captured in the act of egg laying supports the cloaca to egg and nest soil to egg mode for salmonellae infection in the resultant hatchling. Salmonella arizonae and Salmonella serogroups B, C2, and E1 were isolated from the cloaca, colon contents, pond water, and nest soil, and were excreted by hatchlings produced from eggs removed from the soil nests. These same serogroups were isolated from the colon contents of 19 of 37 females tested during the dormant period, suggesting the salmonellae persist in the pond environment in the adult throughout the year.     

The breeding migration of Smooth and Palmate newts (Triturus vulgaris and T. helveticus) at a pond in mid Wales

The breeding population sizes of newts at Llysdinam Pond in 1981 are estimated at around 800 Palmate and 300 Smooth newts. Females were more numerous than males with sex ration of 3.0: 1 for Palamte and 2.6: 1 for Smooth newts. The movement of males of both species into the pond preceded that of females, and female Palmate newts moved out of the pond significantly later than the other classes. The number of females of both species leaving the pond was much less than the number entering, apparently as a result of mortality whilst in the pond. Females leaving the pond were significantly lighter than those entering, and for female Palmate newts the weight loss in only partly attributable to loss of egg load. Temperature and humidity were important factors influencing the number of newts active during the inward migration period, but had little effect on activity during the outward migration.     

Sex allocation in the sexually monomorphic fairy martin

Offspring sex ratios were examined at the population and family level in the sexually monomorphic, socially monogamous fairy martin Petrochelidon ariel at five colony sites over a 4-year period (1993–1996). The sex of 465 nestlings from 169 broods was determined using sex-specific PCR at the CHD locus. In accordance with predicted sex allocation patterns, population sex ratios at hatching and fledging did not differ from parity in any year and the variance in brood sex ratios did not deviate from the binomial distribution. Further, brood sex ratio did not vary with hatching date during the season, brood number, brood size or colony size. The sex ratio of broods with extra-pair young did not differ from those without, while the sex ratio of broods fathered by males that gained extra-pair fertilizations did not differ from broods fathered by other males. Extra-pair chicks were as likely to be male as female. Neither the total number of feeding visits to the brood nor the relative feeding contribution by the sexes varied significantly with brood sex ratio. Brood sex ratios were also unrelated to paternal size, condition and breeding experience or maternal condition and breeding experience. However, contrary to our prediction, brood sex ratio was negatively correlated with maternal size. Generally, these results were consistent with our expectations that brood sex ratios would not vary with environmental factors or parental characteristics, and would not influence the level of parental provisioning. However, the finding that females with longer tarsi produced an excess of daughters is difficult to reconcile with our current understanding of fairy martin life history and breeding ecology.     

Female polyandry and size‐assortative mating in isolated local populations of the Japanese common toad Bufo japonicus

In anurans, female polyandry under male harassment is distributed across taxa because of external aquatic fertilization. According to the sexual selection theory, male–male competition for access to females is affected by the operational sex ratio (OSR) and population density. The Japanese common toad, Bufo japonicus, is widespread in mainland Japan, and like the European common toad, B. bufo, it engages in explosive breeding. In this study, we observed the breeding behaviour of B. japonicus in isolated local populations for over four years in two breeding ponds with different population sizes and densities: large‐low (L‐pond) and small‐high (S‐pond). We analysed the relative polyandry ratio in egg clutches laid by females and estimated the size‐assortative mating pattern to be an indicator of male–male competition in the two ponds. Both ponds tended to exhibit a size‐assortative mating pattern; however, the frequency of polyandry was different in the two ponds (L‐pond = 20% and S‐pond = 90%). Our results showed that polyandry could occur without multiple amplexus with a high population density, i.e. eggs were often fertilized by free‐swimming sperm in the small shallow pond. We propose that high female polyandry ratios without continuous male harassment are generated because of a male‐biased OSR and a high population density in the small pond. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 236–242.     

Gonadal maturation,fecundity, spawning,and timing of reproduction in the mud snail,Cerithidea cingulata,a pest in milkfish ponds in the Philippines

Summary

Gonadal maturation, spawning, fecundity and timing of reproduction of the snail Cerithidea cingulata in a brackish water pond in Molo, Iloilo, Philippines, are described. Snails 4–41 mm in shell length were sampled monthly from May 1997 to May 1998; 25% were <25 mm, 67% were 20–30 mm, and 8% were >30 mm. The sexes are separate and could first be distinguished at 15 mm. Males are aphallic, have narrower shells than females of the same length, and have bright yellow-orange testes overlying the digestive gland deep inside the shell. Females have more robust shells, an ovipositor at the right side of the foot, and yellow-green ovaries overlying the digestive gland. The sex ratio was one male to two females in the pond population studied. Gonadal maturation was monitored by means of gonadosomatic index (GSI, gonad weight as a percent of visceral weight); maturation stages were based on the gonad appearance (immature, developing, mature) and histology (immature, developing, mature, redeveloping). GSI increased with snail size, and reached 16% in a 33-mm female. The smallest mature males and females were 18–19 mm, and most snails >20 mm were mature, spawning, or redeveloping. Histological sections showed all stages of gametogenesis in mature male snails. The oocyte size-frequency distributions in mature females showed mostly mature oocytes and secondary oocytes, but also oogonia and primary oocytes. GSI and the frequency of snails at different maturation stages varied over the year. Both GSI and the frequency of mature snails were highest during the summer months, April to August. Nevertheless, mature snails occurred throughout the whole year, as did mating and egg-laying. Fecundity (= number of oocytes >70 pμ) increased with size in mature females 2041 mm; an average 25-mm female produced about 1,500 oocytes and larger females produced a maximum of about 2,500 oocytes. Eggs strings laid on the pond bottom were 45–75 mm long; an average 64-mm string contained 2,000 eggs 210+20 pm in diameter. The density of eggs strings was highest (80–120/m²) during March-September. Eggs hatched after 6–7 d into planktonic veligers, which in turn settle on the pond bottom 11–12 d later as juveniles. Juveniles 2–6-mm long were most abundant in the pond during August-October.     

Do females adjust the sex of their offspring in relation to the breeding sex ratio?

When the adult sex ratio differs between years in local populations, but still is predictable between adjacent years, it has been proposed that the best strategy would be to bias the offspring sex ratio in favour of the rare sex. We tested this hypothesis using a data set of great reed warbler offspring, sexed by molecular techniques, that were collected over 11 breeding seasons at two adjacent reed marshes. Three important assumptions for this hypothesis are fulfilled in the studied great reed warbler population. First, a substantial proportion of great reed warblers are living in small local populations where sex ratio distortions would be sufficiently large and common. Second, breeding adults and their offspring return to breed in the local population to a high degree. Third, females have a possibility to assess the breeding sex ratio before laying their eggs. At our study site, the breeding sex ratio was positively correlated between successive years. However, contrary to our prediction, female great reed warblers seemed not to adjust their offspring sex ratio in relation to the local breeding sex ratio.     

Breeding and egg survival of a population of a salamander,Hynobius nebulosus tokyoensis Tago

1. A population of a salamander, Hynobius nebulosus tokyoensisTago was studied at Habu, Hinodemachi, in Tokyo during the breeding seasons from 1976 to 1978. Adults appeared in the pond were captured and marked individually by toe-clipping. Egg sacks were counted and tagged with vinyl tapes for the marking.
2. The annual egg production of this population was found to be constant, about 6500 eggs, and total number of breeding adults appeared in the pond also varied little during three years.
3. The appearance of breeding adults in the pond and oviposition seemed to be influenced by the rise of temperature and rainfall.
4. Males appeared earlier in the pond and stayed there for approximately 10 to 20 days. On the other hand, females appeared later in the pond and left there soon after laying eggs.
5. Mean clutch size and body length of breeding adults appeared in the pond showed a tendency to decrease as time proceeded, which seemed to show that the smaller and younger the salamander was, the later it appeared in the pond.
6. Owing to low temperature, the eggs laid earlier took more time to hatch than those laid later.
7. The survival rate of eggs was 67.2% in 1976, 86.4% in 1977 and 81.2% in 1978. The loss rate of eggs laid in the early and late periods of oviposition was higher than that of eggs laid in the middle period.

     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

SEXUAL SELECTION AND FITNESS VARIATION IN A POPULATION OF SMALLMOUTH BASS,MICROPTERUS DOLOMIEUI (PISCES: CENTRARCHIDAE)

Monogamy is often presumed to constrain mating variance and restrict the action of sexual selection. We examined the reproductive patterns of a monogamous population of smallmouth bass (Micropterus dolomieui), and attempted to identify sources of within-season fitness variation among females and known-age males. Many males did not acquire a nest site, and many territorial males were unsuccessful in acquiring a mate. The likelihood that territorial males mated depended on several aspects of nest sites. Mated males of age three were larger than the average size of age-three males in the population. The mean sizes of age-four and age-five mated males were not different from the average of same-age males in the population. Thus, selection resulting from the acquisition of a mate favored large size among only age-three males. Timing of nest construction and breeding among territorial males was negatively related to male size and did not depend on male age after taking male size into account. Indirect evidence (numbers of eggs deposited in nests) suggests that the timing of spawning among females was also negatively related to female size. Fertility selection favored early reproduction within the season by males of all ages, but large male size was favored among only age-four males. The combined early breeding of fecund females and female mate choice of large males may explain the positive correlation between the size of age-four males and the number of eggs acquired. Despite large differences of female fecundity, however, the variance of relative mate number contributed about two times more than the variance of relative fertility among females to the total variance of relative fitness within each sex.     

Arrival to and location of Bufo woodhousei in the breeding pond: effect on the operational sex ratio

Summary Individual male and female Bufo woodhousei spent only two or three nights of the several week breeding season in the pond. In spite of the continual arriving and leaving of individuals the operational sex ratio (the number of females available to mate/number of males attempting to mate) varied little, either temporally across the breeding season or spatially between male groups in the pond. Male body size distributions also varied little between nights and between male groups.     

INEXPLICABLY FEMALE‐BIASED SEX RATIOS IN MELITTOBIA WASPS

The sex ratio behavior of parasitoid wasps in the genus Melittobia is scandalous. In contrast to the prediction of Hamilton's local mate competition theory, and the behavior of numerous other species, their extremely female‐biased sex ratios (1–5% males) change little in response to the number of females that lay eggs on a patch. We examined the mating structure and fitness consequences of adjusting the sex ratio in M. australica and found that (1) the rate of inbreeding did not differ from that expected with random mating within each patch; (2) the fitness of females that produced less female‐biased sex ratios (10 or 20% males) was greater than that of females who produced the sex ratio normally observed in M. australica. These results suggest that neither assortative mating nor asymmetrical competition between males can explain the extreme sex ratios. More generally, the finding that the sex ratios produced by females led to a decrease in their fitness suggests that the existing theory fails to capture a key aspect of the natural history of Melittobia, and emphasizes the importance of examining the fitness consequences of different sex ratio strategies, not only whether observed sex ratios correlate with theoretical predictions.     

Relocation, reproduction and remaining alive in the orb-web spider

When mortality is high, animals run a risk if they wait to accumulate resources for improved reproduction so they may trade-off the time of reproduction with number and size of offspring. Animals may attempt to improve food acquisition by relocation, even in 'sit and wait' predators. We examine these factors in an isolated population of an orb-web spider Zygiella x-notata . The population was monitored for 200 days from first egg laying until all adults had died. Large females produced their first clutch earlier than did small females and there was a positive correlation between female size and the number and size of eggs produced. Many females, presumably without eggs, abandoned their web site and relocated their web position. This is presumed because female Zygiella typically guard their eggs. In total, c . 25% of females reproduced but those that relocated were less likely to do so, and if they did, they produced the clutch at a later date than those that remained. When the date of lay was controlled there was no effect of relocation on egg number but relocated females produced smaller eggs. The data are consistent with the idea that females in resource-poor sites are more likely to relocate. Relocation seems to be a gamble to find a more productive site but one that achieves only a late clutch of small eggs and few achieve that.     

Does breeding population trajectory and age of nesting females influence disparate nestling sex ratios in two populations of Cooper's hawks?

Offspring sex ratios at the termination of parental care should theoretically be skewed toward the less expensive sex, which in most avian species would be females, the smaller gender. Among birds, however, raptors offer an unusual dynamic because they exhibit reversed size dimorphism with females being larger than males. And thus theory would predict a preponderance of male offspring. Results for raptors and birds in general have been varied although population‐level estimates of sex ratios in avian offspring are generally at unity. Adaptive adjustment of sex ratios in avian offspring is difficult to predict perhaps in part due to a lack of life‐history details and short‐term investigations that cannot account for precision or repeatability of sex ratios across time. We conducted a novel comparative study of sex ratios in nestling Cooper's hawks (Accipiter cooperii) in two study populations across breeding generations during 11 years in Wisconsin, 2001–2011. One breeding population recently colonized metropolitan Milwaukee and exhibited rapidly increasing population growth, while the ex‐Milwaukee breeding population was stable. Following life‐history trade‐off theory and our prediction regarding this socially monogamous species in which reversed sexual size dimorphism is extreme, first‐time breeding one‐year‐old, second‐year females in both study populations produced a preponderance of the smaller and cheaper sex, males, whereas ASY (after‐second‐year), ≥2‐year‐old females in Milwaukee produced a nestling sex ratio near unity and predictably therefore a greater proportion of females compared to ASY females in ex‐Milwaukee who produced a preponderance of males. Adjustment of sex ratios in both study populations occurred at conception. Life histories and selective pressures related to breeding population trajectory in two age cohorts of nesting female Cooper's hawk likely vary, and it is possible that these differences influenced the sex ratios we documented for two age cohorts of female Cooper's hawks in Wisconsin.     

Morphometics and Gonadal Development of the Hagfish Eptatretus cirrhatus in New Zealand

Hagfishes have been the target of commercial fisheries in many areas of the world, with the catch processed for leather and for human consumption. A fishery has been operating in New Zealand waters for the last six years, harvesting the bearded hagfish, Eptatretus cirrhatus. The fishery has thus far been unregulated. Based on samples collected dockside over a two-year period, this report expands the morphometric database for this species, provides information on the size and weight of the harvested animals, determines the sizes at the onset of gonadal development and the minimum sizes at sexual maturation for males and females, and indicates that E. cirrhatus, like most other hagfish species, has no specific breeding season. Although females appear in the population at smaller sizes, the sex ratio for mature animals is 1:1 and the sizes of the largest males and females are comparable. The changes observed in sex ratio as a function of TL suggest differences in the timing and rates of gonadal development in females versus males rather than protogyny. Based on the size of the eggs, the number of eggs per female, the proportion of the population that contains large eggs, and the number of postovulatory females, it is clear that E. cirrhatus, like other hagfish species, are potentially vulnerable to overexploitation.     

Oviposition Site Selection and Spatial Predictability of Female White-faced Dragonflies (Leucorrhinia intacta) (Odonata: Libellulidae)

Location of breeding or egg-laying sites may be an important factor underlying social behavior of many organisms. We studied oviposition sites used by a population of Leucorrhinia intacta dragonflies at a small pond near Syracuse, NY, USA. Females preferentially used shallow water as an oviposition habitat. Shallow water increases egg hatching rate through temperature effects on development time and reduces predation on the female. Use of shallow water areas was reduced by locally high densities of territorial males. The same oviposition sites probably maximized the fitness of the male, female, and offspring, meaning that site selection did not necessitate fitness tradeoffs among these classes of individuals. Pond sectors used by ovipositing females were significantly correlated within and between years, but not within days or between consecutive days. Use of pond sectors within and between days was not related to the relative availability of the shallow water habitat, while seasonal use of pond sectors was related positively to availability of shallow water. In spite of the preference by females for ovipositing in shallow water, short-term location of females was not predictable and males could not search predictably good areas during a day. However, across a season, areas with more shallow water available were used for oviposition more than areas with limited substrate.     

Geographical variation in reproductive ageing patterns and life‐history strategy of a short‐lived passerine bird

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle‐aged females (i.e. 2–4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within‐individual increase in timing of breeding and clutch size in early life and a greater within‐individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle‐aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor‐quality members of the pairs, mainly poor‐quality males. Age‐related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life‐history strategies in free‐ranging animals from more than a single population can throw new light on life‐history theory, population dynamics and evolutionary studies of senescence.     

Population variation and ecological correlates of tychoparthenogenesis in the mayfly, Stenonema femoratum

Species in which both sex and parthenogenesis co‐occur are extremely valuable for investigating ecological conditions favouring sex. Tychoparthenogenesis is a breeding system characterized by hatching of a small proportion of unfertilized eggs (typically < 10%) from females of sexually reproducing species. With tychoparthenogenesis, both sexual and parthenogenetic reproduction co‐occur within the same population. To identify ecological conditions that may favour this breeding system, I quantified population variation in females’ capacity for tychoparthenogenesis and investigated biotic and abiotic correlates of tychoparthenogenesis. I estimated tychoparthenognetic capacity (proportion of unfertilized eggs hatching) for females from 12 Missouri populations of the mayfly, Stenonema femoratum (Ephemeroptera: Heptageniidae), across three different habitat types – temporary streams, permanent streams and lakes. Tychoparthenogenetic capacity, measured as the population mean hatch success of unfertilized eggs, ranged from 3.8 to 10.7%. Tychoparthenogenetic capacity varied among habitats in 1996, but not in 1997. In 1996, temporary streams showed hatch success of unfertilized eggs twice that of permanent streams and lakes. Tychoparthenogenetic capacity also varied among sampling dates within years. Temporary streams also showed extremely low nymph densities compared to the other two habitats. However, habitats did not differ in adult density. Furthermore, in all populations nymphs showed significantly female‐biased sex ratios. In contrast, adult sex ratios were equal or slightly male biased. Tychoparthenogenetic capacity was negatively correlated with nymph density in 1996, but not in 1997, suggesting possible reproductive assurance in some years. Adult densities also suggested that there may be certain times of year when tychoparthenogenesis may provide benefits of reproductive assurance. Although habitats differed significantly in their abiotic characteristics, tychoparthenogenetic capacity was correlated significantly with water temperature only. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75 , 101–123.