Rapid growth and large body size in annual fish populations are compromised by density-dependent regulation

We tested the effect of population density on maximum body size in three sympatric species of annual killifishes Nothobranchius spp. from African ephemeral pools. We found a clear negative effect of population density on body size, limiting their capacity for extremely fast development and rapid growth. This suggests that density-dependent population regulation and the ephemeral character of their habitat impose contrasting selective pressures on the life history of annual killifishes.     

Reproductive strategies in rain-pool dwellers and the model freshwater insect

Reproductive strategies of principal rain-pool midges, Chironomus imicola and Chironomus pulcher are investigated in field and laboratory. The two species are able to lay at least one full clutch without access to food in the adult stage. Since females are also capable of flying for extended periods without feeding, they carry out both adult functions (i.e. dispersal and reproduction), without taking energy from the terrestrial environment.We argue that independance of the terrestrial environment is adaptively appropriate to animals able to exploit a larval habitat rich in food and low in interspecific competition. However, there are costs to this strategy because freshwaters, and notably rain-pools, are unpredictable habitats inclined to dry up periodically. These species must, therefore, constantly colonise new pools by laying eggs. We show that adult females can resort to feeding and that if they do so the energy acquired goes to the production of further eggs with a consequent increase in colonizing ability.As a test of this hypothesis that there is a relationship between duration of habitat and the number of eggs layed, we have examined a third rain-pool dwelling chironomid, Polypedilum vanderplanki. This species is unique among insects in that larvae are able to survive desiccation of their tissues. For them there is thus not the same incentive to leave before the home pool dries. So, P. vanderplanki does not need to be a colonizer. Observations confirm this view since P. vanderplanki adult females, even when fed, lay no more than one clutch of eggs. Consequently, reproduction and feeding in the adult female are related to the way in which duration of the habitat is perceived. We suggest that, measured against P. vanderplanki, Chironomus species provide a useful model of the ideal freshwater animal.     

Hatching phenology,life history and egg bank size of fairy shrimp Branchipodopsis spp. (Branchiopoda,Crustacea) in relation to the ephemerality of their rock pool habitat

In temporary aquatic habitats, permanence and the severe disturbance associated with desiccation are strong selective agents expected to lead to differentiation in life history strategies in populations experiencing different disturbance regimes. Besides optimal timing of hatching of dormant life stages, maturation and reproduction, pool inhabitants also benefit from the acquisition of reliable cues for the quality of the ambient environment. We investigated whether hatching patterns, life history characteristics and egg bank size of Branchipodopsis fairy shrimp (Branchiopoda, Anostraca) inhabiting a cluster of temporary rock pools in South Africa reflect variation in habitat stability and hatching cues. Long-term hydrological variation was used to select pools along a gradient of habitat stability. Initial conductivity was a good indicator for the length of inundations. No hatching occurred under elevated conductivities, which may present a mechanism to avoid abortive hatching. Egg bank size was unaffected by habitat size or habitat stability but instead was related to cover by a protective sheet of dry aquatic vegetation, which presumably counteracts egg bank erosion by wind when pools are dry. Life history but not hatching phenology reflected some aspects of habitat stability. Fairy shrimp populations in ephemeral pools started reproduction earlier than populations in more stable habitats. Additional common garden or transplant experiments, however, will be required to assess the relative importance of environmental and genetic components in explaining the observed variation and acquire more insight into the trade-offs that lie at the base of the evolution of life history strategies along the pond permanence gradient.     

Host range testing of a prospective classical biological control agent against cabbage maggot,Delia radicum,in Canada

The introduction of a European natural enemy, Aleochara bipustulata L. (Coleoptera: Staphylinidae), is being considered for control of cabbage maggot, Delia radicum (L.) (Diptera: Anthomyiidae) in canola in Canada, and the host specificity of this pupal parasitoid must first be studied. Contemporary guidelines were used to select 18 species of Diptera to represent non-target species taxonomically related or ecologically similar to reported hosts of A. bipustulata, or beneficial species. No-choice tests were used to determine which of the 18 species are within A. bipustulata’s fundamental host range, and whether puparial structure or mass or duration of pupal development influence their acceptability and suitability as hosts. Five species were consistently suitable as hosts, and these were either relatively small or were taxonomically closely related to the target host. The probability that a puparium would be entered by a parasitoid larva was greatest for small puparia, but was unaffected by pupal duration. The probability of completing parasitoid development once a puparium was entered was influenced by both puparial mass and duration of pupal development. Pitfall traps to assess habitat associations caught A. bipustulata adults in a variety of crop habitats but none were caught in forests. Host range and habitat data are used to argue that there is little risk of parasitism to beneficial taxa. Non-target species taxonomically closely related to the target D. radicum or with small puparia may fall within the fundamental host range of A. bipustulata. However, risk to many of these species may be minimal because of the habitat preferences of the parasitoid and its cues for host-finding and recognition.     

Successional phases and species replacements in freshwater rock pools: towards a biological definition of ephemeral systems

1. In temporary aquatic habitats, time is probably the dominant environmental factor affecting community composition, mainly by setting constraints on colonization success and the replacement of taxa over time. The mechanism and effect of a decreasing inundation period on community development, mostly in terms of truncation, are still poorly documented. The permanent and ephemeral components of temporary communities are expected to be differently influenced by the degree of persistence of the habitat. 2. To study the effect of time on invertebrate community assembly and dynamics in a short duration type of temporary aquatic habitat, we monitored 16 ephemeral rock pools which persisted from less than a week to about 1 month at two rock pool sites in semi‐arid south‐eastern Botswana. Data were collected every 2 days during a full inundation cycle. 3. All communities were initially assembled by permanent residents recolonizing the habitat from egg banks and were later joined by actively dispersing ephemeral taxa. Species replacements only occurred in two pools. Concurrent with a decrease in the densities of Branchipodopsis wolfi, population sizes of Leberis sp. and Culicidae (Aedes sp. and Anopheles sp.) increased in these pools. Although it was possible to distinguish two successive phases at one rock pool site, community assembly was generally a gradual process determined by dispersal strategies of the inhabitants. Additional rains after initial filling triggered dispersal by ephemeral taxa, mainly Micronecta youngiana and Hydroglyphus infirmus, and positively influenced colonization success. 4. Decreasing persistence shortens community development down to a critical point below which lack of time eliminates the possibility of species replacement. Based on these findings, we define ephemeral waters as aquatic habitats lacking species replacements. Other temporary water types have a relatively longer persistence, permitting successional replacement of species.     

Interaction between insect larvae and tadpoles in tropical rain pools

Abstract. 1. Ephemeral rain pools on rock surfaces are common in Africa and are inhabited by dense populations of aquatic dipteran larvae. About 30% of the pools also support large numbers of tadpoles of the frog Ptychadena anchietae .
2. Experiments reveal that the presence of tadpoles suppress eclosion of the rock pool dwelling midge Chironomus imicola .
3. However, the presence of tadpoles also shortens the larval life-span of C. imicola by speeding up growth rates. Since the larval stages must be completed before the pool dries tadpoles may help 'fine-tune' the dipteran to the rock pool habitat.
4. A change in the diet of Cimicola larvae is associated with the accelerated development. Grazing by tadpoles results in algae, growing on the water surface, reaching the mud in tadpole faeces. These algae represent a high protein and energy food not otherwise accessible to mud dwelling dipteran larvae.
5. When pools dry tadpoles are killed, but a second species of dipteran Dasyhelea thompsoni have larvae able to survive to dry phase in situ . These larvae are scavengers so that on reflooding dead tadpoles are available as food. Experiments show that the presence of tadpoles again increase larval growth rates.
6. Both alive and dead tadpoles therefore are responsible for increasing the quality and quantity of food available to dipteran larvae and are thus among the mechanisms ensuring an abundance of food for rock pool dwelling insect larvae.     

Breeding habitat selection and larval performance of two anurans in freshwater rock-pools

I studied breeding habitat choice of common toad Bufo bufo and common frog Rana temporaria in 221 freshwater rock-pools on 15 small islands in the southwestern coast of Finland. I monitored tadpole growth rate in a number of rock-pools differing in size, competitors and predator regime. Furthermore, I carried out a short-term field experiment to investigate the effects of different predators on larval survival. Rana bred in pools of virtually all sizes, although the very smallest ones were usually avoided. A logistic regression analysis showed that the amount of aquatic macrophyte vegetation and pool surface area were the most important pool characters for Rana. Bufo preferred the largest pools, and pool surface area was by far the most important habitat variable. Rana avoided pools with the lowest pH. but was more common than expected at pools with pH around 6. In Bufo such an effect was not found, but this may be due to relatively low number of pools occupied by Bufo. In Rana there was a positive correlation between pool area and tadpole body length in early June, but two weeks later this relationship was not significant. In Bufo tadpole body size was not significantly affected by pool area. Presence of predators or presence of the other tadpole species did not affect growth or tadpole body size in either Rana or Bufo. A short-term experiment in wading pools indicated that both species suffered heavy mortality in the presence of a diving beetle larva. In the presence of predatory fish, mortality of Bufo did not differ from controls, whereas mortality was high among Rana tadpoles. It appears that although predators prey heavily on Rana tadpoles, they do not exclude Rana from larger rock-pools. However, it is not evident from this study why Bufo avoids smaller pools.     

Ochlerotatus albifasciatus in rain pools of Buenos Aires: seasonal dynamics and relation to environmental variables

The immature stages of Ochlerotatus albifasciatus develop in temporary pools. The present study aims at evaluating the seasonal dynamics of the aquatic stages of this mosquito, also analyzing the relationship among their presence and breeding success to some relevant climatic and environmental variables in the ephemeral rain pools of an urban park. Nineteen cohorts of O. albifasciatus that developed synchronously after rain events were recorded in all seasons. The proportions of mosquito-positive pools were significantly higher during the fall-winter period than in the spring-summer months (p < 0.001). The presence of this mosquito species was positively related to the amount of rain (p < 0.001), whereas negatively correlated to air temperature (p < 0.05) within a 5.2 to 29.7 degrees C range. The distribution of the number of cohorts per pool throughout the year was grouped (variance/mean: 3.96), indicating that these habitats were not equally suitable as breeding sites. The immature stages of O. albifasciatus were detected in pools belonging to all of the categories of surface area, depth, duration, vegetation cover, and insolation. However, the proportion of pools where immature mosquitoes were detected was positively and significantly related to surface, depth, duration, and vegetation cover. On the other hand, the proportion of mosquito-positive pools was higher at an intermediate insolation degree. Our results suggest that although preimaginal stages were present in all seasons, high temperatures may be unfavorable to larval development, and substrate vegetation may regulate water temperature. The positive relationship between the proportion of mosquito-positive pools and pool size and duration might reflect a strategy of O. albifasciatus to accomplish immature development.     

Rain-pools on peat moorland as island habitats for midge larvae

A common aquatic habitat on peat moorland is a pool, often only a few centimetres deep. Many have a shorter average duration than that of the inhabitants but they are, nonetheless, inhabited by the larval stages of several chironomid species at moderately high densities. Because of their durational characteristics, these pools might be expected to have an endemic fauna, as is the case in similar ephemeral island habitats elsewhere. However, we find no such fauna. Indeed, we find no qualitative difference between them and permanent pools of the same substratum.We identify three reasons for the apparent absence of endemism. First, the midge fauna is able to survive dry periods in the moist mud, so that even the smallest pools are effectively permanent from the adaptive point of view. Second, although seasonal freezing eliminates the fauna of smaller pools, larvae are able to complete their development in the preceding summer. Third, nearby permanent pools which do not dry and in which the mud does not freeze, may act as refugia for re-invasion.     

Large and least isolated fragments preserve habitat specialist spiders best in dry sandy grasslands in Hungary

The role of fragment size, isolation and habitat diversity in the conservation of spider assemblages living in fragmented landscape were studied in dry sandy grasslands (East Hungary, Nyírség). Spiders were collected using pitfall traps at eight dry grassland fragments from 2001 to 2009 from March to October fortnightly. We tested the rules of island biogeography, which suggest that the species richness increases with the size and decreases with the isolation of fragments. The habitat diversity is an important factor for species richness, since large areas usually have more habitats; therefore, the number of species may be higher in these areas. During the 9-year study period, altogether 10,544 individuals belonging to 106 species were collected. Contradicting the classical theory, we found a significant negative relationship between the total number of spider species and the grassland size, while the ratio of sandy grassland specialist spider species increased with fragment size. The relationship between the ratio of generalist species and the fragment size was not significant. The overall species richness and the isolation of studied grasslands did not show a significant relationship. The ratio of sandy grassland specialist species decreased, while the ratio of generalist species increased with the increasing of isolation. The habitat diversity did not show any effect on spider species richness. We concluded that to conserve the habitat specialist species it is recommended to preserve the large and least isolated grassland fragments, furthermore to increase the size of small fragments with the restoration of the adjacent croplands.     

Body mass, habitat generality, and avian community composition in forest remnants

Aim Many theories of biodiversity and biogeography assume that species respond equally to variability in habitat area and isolation. This assumption does not allow for differential responses due to interspecific competition or other mechanisms, and therefore does not allow community composition to be predicted. As body size is relevant to area requirements and interspecific dominance, a natural experiment was conducted to quantify the differential responses of avian species abundance to variability in remnant area, isolation and forest cover based on average species body mass. Location Deciduous forest remnants of varying area and isolation throughout the State of Delaware, USA. Methods Forest remnants within stratified area and isolation classes were randomly selected for breeding bird surveys; total forest cover (ha) within 2 km of each survey point was subsequently determined as a covariate. Surveys were conducted within 100–150 m from the edge of each remnant and detected bird species were divided into five classes based on a log₂‐transformation of body mass (very small, small, medium, large and very large). Assuming a negative binomial distribution, the abundance of detected individuals in each mass class was analysed using generalized linear models with remnant area, isolation, local forest cover and two‐way interactions specified as independent variables. The same analyses were conducted for individual species where sample size allowed. Results Very small, small and very large bodied species decreased in abundance with decreasing local forest cover and remnant area and with increasing remnant isolation, while large species increased in abundance. Medium‐sized species decreased in abundance with increasing forest cover, did not respond to remnant isolation and showed a concave, curvilinear response to increasing remnant area. Large and medium‐sized species were the most abundant birds in small, isolated remnants despite occurring in the largest remnants with the more abundant very small and small species, suggesting that communities are not randomly organized. Main conclusions Regardless of presumed habitat associations, large and medium‐sized species are of the appropriate size to be dominant competitors when forest resources are limiting, and thus may be considered ‘generalists’. The smallest species may be excluded entirely from small, isolated remnants even though such remnants meet their ecological needs; the needs of very large species are not met in small remnants. The applicability of biodiversity theories to community composition, species abundance and, by extension, to conservation, can be improved by incorporating differential responses based on body mass into their assumptions.     

Overcoming barriers to active interventions for genetic diversity

As a result of processes such as habitat loss and overharvest, many species persist in small, isolated populations that experience reduced fitness, decreased evolutionary potential, and increased extinction risk. The goal of species conservation is to restore genetic diversity and adaptive potential caused by isolation and small population size. For populations trapped in an extinction vortex, habitat protection may be inadequate for successful conservation. Alternative actions such as deliberate admixture by introducing individuals from related subspecies may be necessary to recover population fitness. While there is precedent for such actions, admixture temporarily disrupts the taxonomic integrity of a species. Concerns for the taxonomic integrity or “naturalness” of a species may prevent the use of active interventions that involve admixture and transient hybrid gene pools even though extinction may be imminent. We explore the cultural barriers to using tools such as genetic rescue and make suggestions for overcoming those barriers. We focus mainly on examples from animals, but the same evolutionary processes are ongoing in other life forms and are subject to the same cultural barriers.     

Life history consequences of feeding versus non-feeding in a facultatively non-feeding toad larva

Summary Bufo periglenes, a toad endemic to montane Costa Rica, produces an unusually small clutch of large, yolk-rich eggs. The toads breed in small ephemeral pools that are unpredictable in duration and may be low in food availability. Two congeners, Bufo coniferus and Bufo marinus, occur nearby, breed in more permanent bodies of water that offer more food, and exhibit the typical toad pattern of large clutches of small eggs. Tadpoles of all three species feed on detritus and suspended organic material. By raising tadpoles of the three species individually with and without food I investigated the relationship between egg size (yolk provision) and tadpole survival. All of the unfed B. coniferus and B. marinus tadpoles grew little and died soon after developing to the hindlimb bud stage. On the other hand, all of the unfed B. periglenes tadpoles metamorphosed successfully, demonstrating that the tadpoles are facultatively non-feeding; developmental time from hatching to metamorphosis was significantly shorter for unfed tadpoles than for fed tadpoles, but fed individuals were significantly larger at transformation. Faster developmental rate and larger body size at transformation are both advantageous for frogs and toads, but cannot be attained simultaneously. Large egg size may afford flexibility in unpredictable environments. In pools where food is available, tadpoles presumably eat, take longer to metamorphose, but are larger at transformation than tadpoles developing in nutrient-poor sites. Small body size at transformation (a consequence of not eating) has potential costs, but the large quantity of yolk provided by a large egg enhances the probability of metamorphosis in food-limited environments.     

Scale-dependence and mechanisms of dispersal in freshwater zooplankton

Communities of organisms form as a result of both interspecific and abiotic interactions within local habitat patches and dispersal among patches in a region. Local processes are expected to play a dominant role when dispersal occurs much more often than extinction. We performed two field experiments to examine rates and mechanisms of dispersal in freshwater pond zooplankton communities. First, we tested the effect of distance from a source on the rate of colonization of artificial habitat by placing wading pools at 5, 10, 30 and 60 m from two natural fishless ponds and observing the succession of zooplankton. Seventy-eight percent of the species in the source ponds that were capable of living in the pools colonized at least once during the experiment. A new species was found in the pools on average once every four days, suggesting that colonization events occur on the order of days to weeks for many species. Colonization rates declined further from the source at one pond but not the other, and the effect of distance was relatively weak at both ponds. This suggests that many species disperse broadly over short distances. The second experiment tested the role of animal vectors for zooplankton dispersal by restricting access to the pools. Eight treatments were imposed that excluded potential animal vectors along a body size gradient from large mammals to small insects. While the treatments affected zooplankton colonization, many species invaded even when all animals larger than 1 mm were excluded. Animal vectors may therefore be less important for dispersal than wind. Our results suggest that zooplankton are highly effective dispersers over short distances, and can disperse via several mechanisms. Local interactions should therefore play a dominant role in structuring these communities at small regional scales.     

Local structuring factors of invertebrate communities in ephemeral freshwater rock pools and the influence of more permanent water bodies in the region

We used three isolated clusters of small ephemeral rock pools on a sandstone flat in Utah to test the importance of local structuring processes on aquatic invertebrate communities. In the three clusters we characterized all ephemeral rock pools (total: 27) for their morphometry, and monitored their water quality, hydrology and community assemblage during a full hydrocycle. In each cluster we also sampled a set of more permanent interconnected freshwater systems positioned in a wash, draining the water from each cluster of rock pools. This design allowed additional testing for the potential role of more permanent water bodies in the region as source populations for the active dispersers and the effect on the community structure in the rock pools. Species richness and community composition in the rock pools correlated with level of permanence and the ammonia concentration. The length of the rock pool inundation cycle shaped community structure, most probably by inhibiting colonization by some taxa (e.g. tadpoles and insect larvae) through developmental constraints. The gradient in ammonia concentrations probably reflects differences in primary production. The more permanent water bodies in each wash differed both environmentally and in community composition from the connected set of rock pools. A limited set of active dispersers was observed in the rock pools. Our findings indicate that aquatic invertebrate communities in the ephemeral rock pools are mainly structured through habitat permanence, possibly linked with biotic interactions and primary production. Handling editor: S. I. Dodson     

Survival strategies in tropical rain pools

Summary Adaptations of three larval diptera in central Africa, each inhabiting rain-filled rock pools of characteristic duration, have been studied by field observation and experiment.The duration of the pool is important in determining which of the three species is present. Large pools, lasting a few weeks after each rain, favour Chironomus imicola which has a larval life span of as little as twelve days. This species relies on egg laying female flies to re-invade newly filled pools. In cases where the pool lasts for a shorter time than the minimum larval life span a mechanism is needed to survive dry periods in situ. Such pools are inhabited by larvae of Polypedilum vanderplanki and Dasyhelea thompsoni. Larvae of P. vanderplanki are poor at invading newly flooded pools and are also poor at competing with other species. However they are able to tolerate virtually a complete loss of body water and are therefore able to survive drought in the dry mud. They are consequently always the first there after refilling, a factor that provides them with sufficient advantage to eventually become established in very small pools. D. thompsoni larvae, not quite so good at surviving drought, but better at invading, occupy pools of intermediate size. They probably inhabit a larger number of pools than either of the other two species.A hazard associated with surviving dry phases in situ is vulverability to attack by terrestrial scavengers. However the activity of scavengers such as pheidolid ants, although conspicuous, is confined to periods when standing water has evaporated but the mud is still moist. Once dry, predation stops. Losses from this cause are therefore likely to be associated with periodic dry phases throughout the rainy season rather than with the dry season itself.     

Environmental filtering predicts plant‐community trait distribution and diversity: Kettle holes as models of meta‐community systems

• Meta‐communities of habitat islands may be essential to maintain biodiversity in anthropogenic landscapes allowing rescue effects in local habitat patches. To understand the species‐assembly mechanisms and dynamics of such ecosystems, it is important to test how local plant‐community diversity and composition is affected by spatial isolation and hence by dispersal limitation and local environmental conditions acting as filters for local species sorting.We used a system of 46 small wetlands (kettle holes)—natural small‐scale freshwater habitats rarely considered in nature conservation policies—embedded in an intensively managed agricultural matrix in northern Germany. We compared two types of kettle holes with distinct topographies (flat‐sloped, ephemeral, frequently plowed kettle holes vs. steep‐sloped, more permanent ones) and determined 254 vascular plant species within these ecosystems, as well as plant functional traits and nearest neighbor distances to other kettle holes.Differences in alpha and beta diversity between steep permanent compared with ephemeral flat kettle holes were mainly explained by species sorting and niche processes and mass effect processes in ephemeral flat kettle holes. The plant‐community composition as well as the community trait distribution in terms of life span, breeding system, dispersal ability, and longevity of seed banks significantly differed between the two habitat types. Flat ephemeral kettle holes held a higher percentage of non‐perennial plants with a more persistent seed bank, less obligate outbreeders and more species with seed dispersal abilities via animal vectors compared with steep‐sloped, more permanent kettle holes that had a higher percentage of wind‐dispersed species. In the flat kettle holes, plant‐species richness was negatively correlated with the degree of isolation, whereas no such pattern was found for the permanent kettle holes.Synthesis: Environment acts as filter shaping plant diversity (alpha and beta) and plant‐community trait distribution between steep permanent compared with ephemeral flat kettle holes supporting species sorting and niche mechanisms as expected, but we identified a mass effect in ephemeral kettle holes only. Flat ephemeral kettle holes can be regarded as meta‐ecosystems that strongly depend on seed dispersal and recruitment from a seed bank, whereas neighboring permanent kettle holes have a more stable local species diversity.

     

Desiccation risk favours prevalence and diversity of tardigrade communities and influences their trophic structure in alpine ephemeral rock pools

Hydrobiologia - Rock pools are ephemeral freshwater habitats characterized by their small size, well-defined boundaries, and periodic desiccation, making them ideal model systems to answer numerous...     

Water temperature fluctuations and territoriality in the intertidal zone: two possible explanations for the elevational distribution of body size in Graus nigra

On the central coast of Chile, distribution of body size in Graus nigra varied with tidal pool height. With the objective of determining whether environmental temperature is one of the possible causes which explains the observed distribution pattern, two behavioural responses were analysed during an experimental period of increasing water temperature: number of opercular movements (an indirect measure of energy expenditure) and activity levels. The interactions of temperature × time and body size × time had a significant effect on the number of opercular movements. At low temperatures (13–15° C), large fish reached a maximum number of opercular movements, while small fish reached a maximum only at high temperatures (23–25° C). The interaction temperature × time had a significant effect on activity levels of different body sizes. In general, large fish appeared to be less active than small fish, however, at very high temperatures (24–26° C) all individuals increased their activity levels. These data indicate that small fish are acclimatized to live in a wider range of temperatures (13–23° C), and, for fish of all body sizes, the highest temperatures (23–26° C) probably constitute a suboptimal microhabitat. Strong territoriality was observed, with large individuals displacing smaller individuals. These data suggest that temperature is an important factor in explaining why large individuals are not present in high tidal pools (high temperatures), whereas territoriality explains why small individuals are not in low tidal pools (habitat of large individuals).