Can two species of midge coexist in a single puddle of rain-water?

Larvae of two midge species, Chironomus pulcher and Chironomus imicola normally occur in different puddles of rain-water; C. pulcher in shaded and C. imicola in sunny ones. When the sun/shade signal is unclear, both species can coexist for most of each rainy season. Yet, in the laboratory, C. pulcher shows total competitive exclusion of C. imicola. The puzzle is resolved by experiments and observations which suggest that some naturally occurring puddles possess characteristics not present in laboratory pools. These include a spatial refuge (deep water) and separate shaded and sunny parts. Both variables are normally found only in larger puddles, so coexistence is a function of patch size. On its own, each variable is a necessary but not a sufficient condition for coexistence.     

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.     

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.     

Are females selected against in ephemeral habitats? The case of a holometabolous insect (Chironomus pulcher)

Adults of the rain-pool dwelling midge Chironomus pulcher are sexually dimorphic with females larger than males. This difference is achieved by females remaining in the growing larval stage for longer. When food is scarce, as under crowded conditions, the average size of both sexes is reduced, but the difference between them is maintained. However, more females perish, presumably because they are unable to achieve a minimum female size. Under conditions of crowding, there is thus a switch in favour of males. Since crowding due to a diminishing habitat is a frequent event in rain-pools, males seem fitter than females in C. pulcher. This paradox is discussed.     

Phylogeography,population genetics and conservation of the European red deer Cervus elaphus

• 1 During the Last Glacial Maximum, European red deer Cervus elaphus occurred in refugia in Iberia/southern France, Italy, the Balkans and the Carpathians. Most of Europe, including large parts of the east and north‐east, is now inhabited by red deer from the western lineage. The eastern lineage is largely confined to south‐eastern Europe; a third lineage comprises Sardo‐Corsican and Barbary red deer.
• 2 Sardo‐Corsican, Barbary and Mesola red deer are genetically unique units. They exhibit low levels of genetic diversity and deserve particular protection, since conservation strategies should target genetic information.
• 3 Hybridization between sika Cervus nippon and red deer occurs rarely, but may lead to extensive introgression, particularly in parts of the British Isles. Further expansion of both species may lead to increased hybridization in continental Europe.
• 4 Although hunting has an impact on red deer gene pools, the main threat today is habitat fragmentation in human‐dominated landscapes. The resulting increase in genetic drift and inbreeding reduces variability in isolated populations and may lead to inbreeding depression. To support vital meta‐populations, migration corridors should be established.

     

Life in the puddle: behavioural and life-cycle adaptations in the Diptera of tropical rain pools

Puddles of rain water on the surfaces of rock exposures are a little known but very common habitat for freshwater-dwelling animals. In Africa, these are inhabited by the larvae of two taxa of fly unique to these pools. One of these includes species able to survive dry periods in situ; the other includes species that must reach adulthood and migrate to survive periods when the pool is dry. Hence, the opportunity exists for a comparative study of adaptation among these species. Since puddles are small, our principal method in the study of adaptation has been the experimental manipulation of puddles and their faunas in the wild. Using this method we were able to identify the spatial consistency of pools and their unpredictable duration during the rainy season as the main selective pressure shaping adaptation. Adaptations include diapause and adaptive adjustment of the life cycle. It is the second of these that provides the focus of our interest here.     

Modelling the longitudinal distribution,abundance, and habitat use of the giant freshwater shrimp (Macrobrachium spinipes) in a large intermittent,tropical Australian river to inform water resource policy

1. Water development threatens rivers and their biodiversity. Amphidromous shrimp are particularly vulnerable as they require migration between freshwater and estuaries to complete their life cycle. The Fitzroy River is a large tropical intermittent river undergoing water development that is home to the amphidromous shrimp Macrobrachium spinipes (cherabin), yet little is known about its habitat use and flow-ecology making it difficult to inform sustainable water-take.
2. We investigated habitat associations, distributional patterns suggestive of amphidromy, and the influence of water availability by sampling main channel and floodplain pools along a 350-km river length during 2 contrasting flow years. Applying a size-specific abundance model, we estimated abundance per size class, site, and year. We then predicted abundance at the landscape scale with remotely sensed water to reveal the impact of water availability on the meta-population.
3. Our model revealed that juveniles were in greatest abundance in downstream main channel pools, whereas adults were in greatest abundance in upstream floodplain pools. Abundance varied by year with lower numbers predicted in the low-flow year. Longitudinal and habitat patterns remained when our pool-level results were scaled to the landscape, and the positive relationship of abundance to wet-season flow was strengthened. The predominance of smaller cherabin in the lower reaches of the river provides indirect support for an estuarine nursery and amphidromous life history; however, small individuals observed in landlocked pools, during late dry season suggests possible within-river recruitment.
4. The importance of water development policies that protect wet-season flow and passage along the Fitzroy River is supported by this work. These types of policies are likely to be important for this and other amphidromous shrimp species across Australia, Southeast Asia and further afield. Further research detailing the species life history and describing flow–recruitment relationships will be important contributions to understanding this important taxonomic group and refining policies for current and future water resource development.

     

Do pools impede drift dispersal by stream insects?

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Effects of dams and their environmental impacts on the genetic diversity and connectivity of freshwater mussel populations in Poyang Lake Basin,China

1. Habitat fragmentation is one of the main threats to biodiversity. Barriers to dispersal caused by anthropogenic habitat alteration may affect phylogeographic patterns in freshwater mussels. Knowledge of the phylogenetic and phylogeographic patterns of unionoids is vital to inform protection of their biodiversity.
2. Here, we assessed influences of dams and their environmental effects on the genetic diversity and population connectivity of a broadly distributed freshwater mussel, Nodularia douglasiae, in Poyang Lake Basin.
3. The results showed high genetic diversity in areas without dams and low genetic diversity in areas with dams. High genetic differentiation and low gene flow were found among the 11 populations. Genetic variation was significantly correlated with dissolved oxygen levels.
4. The observation of low genetic diversity in populations separated by dams indicated that those populations were subjected to genetic erosion and demographic decline because they are disconnected from other populations with higher diversity. High genetic differentiation and low gene flow among the 11 populations could be correlated with anthropogenic habitat alteration.
5. These results indicated that anthropogenic habitat alterations have led to the decline in freshwater mussel diversity. Therefore, we recommend maintaining favourable habitat conditions and connectivity of rivers or lakes, and strengthening study of life histories with host-test experiments to identify potential host fish species to strengthen the knowledge base underpinning freshwater mussel conservation.

     

Habitat selection as a source of inter‐specific differences in recruitment of two diadromous fish species

• 1 For aquatic species with highly dispersive offspring, the addition of new individuals into an area (recruitment) is a key process in determining local population size so understanding the causes of recruitment variability is critical. While three general causative mechanisms have been identified (the supply of individuals, habitat selection and mortality), we have a limited understanding of how variation in each is generated, and the consequences this may have for the spatial and temporal distribution of recruits.
• 2 We examined whether active habitat selection during settlement could be the cause of variability in populations of two diadromous fish species using a field survey and laboratory‐based choice experiments. If larval behaviour is important, we predicted there would be inter‐specific differences in abundance between sites during the survey, and that larvae would prefer water collected from sites with higher conspecific abundances during the experiments.
• 3 During the field survey, significant differences were detected between two rivers (the Cumberland and Grey), with one species (Galaxias maculatus) found in higher abundances at one site (the Cumberland River) while comparable numbers of a closely related species (Galaxias brevipinnis) were caught at both sites. Laboratory choice experiments were conducted to determine whether larval preferences during settlement could be the cause of these differences. G. maculatus larvae showed a preference for freshwater over saltwater, indicating that the fish may be responding to reduced salinities around river mouths during settlement. The results of a second experiment were consistent with the notion that larval preferences could be the mechanism driving differences in the populations of the two rivers, with G. maculatus preferring water collected from the Cumberland River while G. brevipinnis did not prefer water from either river.
• 4 These results demonstrate that active habitat selection may be important in establishing spatial patterns of larvae at settlement, and that multiple cues are likely to be involved. This study also demonstrates that the behaviours exhibited by individuals can strongly influence the structure and dynamics of populations of aquatic species with complex life cycles.

     

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.     

Effects of extreme low flows on freshwater shrimps in a perennial tropical stream

• 1 Long-term data on rainfall suggests that perennial rainforest streams rarely are subject to drying of riffles or pools in the wet, non-seasonal Caribbean climate of Puerto Rico. Unusually low rainfall in 1994 caused some headwater riffles to dry out completely, resulting in isolated pools, reduced pool volumes and loss of access to microhabitats by benthic invertebrates.
• 2 From 1992 to 1998, shrimp populations were sampled bimonthly using baited traps in six pools along 1200 m (from 305 to 480 m in altitude) of Quebrada Prieta, a second-order headwater stream in the Luquillo Experimental Forest (Caribbean National Forest).
• 3 Following contraction of the smaller and shallower pools in the most upstream section of the stream, mean densities of the dominant shrimp (Atya lanipes) increased from 22 to 75 shrimp m⁻² of pool area during the 1994 drought year. A second common species (Xiphocaris elongata) increased from 5 to 14 shrimp m⁻². A smaller percentage of adults of both species was gravid during the drought.
• 4 Following the 1994 drought (1995–1998), densities of both shrimp species and reproductive activity of Atya returned to predrought (1990–1993) levels. However, the reproductive activity of Xiphocaris remained lower than in the predrought period.
• 5 It is suggested that prolonged droughts, even in tropical rainforest biomes, may significantly alter aquatic communities through localised crowding effects resulting from habitat contraction, and lead to prolonged decreases in reproductive output. Consequently, major alterations in aquatic populations and communities would be predicted by current climate change scenarios of decreased total rainfall and increased variability.

     

Impact of river dynamics on the genetic variation of Gypsophila repens (Caryophyllaceae): a comparison of heath forest and more dynamic gravel bank populations along an alpine river

• Alpine rivers are, despite anthropogenic water flow regulation, still often highly dynamic ecosystems. Plant species occurring along these rivers are subject to ecological disturbance, mainly caused by seasonal flooding. Gypsophila repens typically grows at higher altitudes in the Alps, but also occurs at lower altitudes on gravel banks directly along the river and in heath forests at larger distances from the river. Populations on gravel banks are considered non‐permanent and it is assumed that new individuals originate from seed periodically washed down from higher altitudes. Populations in heath forests are, in contrast, permanent and not regularly provided with seeds from higher altitudes through flooding. If the genetic structure of this plant species is strongly affected by gene flow via seed dispersal, then higher levels of genetic diversity in populations but less differentiation among populations on gravel banks than in heath forests can be expected.
• In this study, we analysed genetic diversity within and differentiation among 15 populations of G. repens from gravel banks and heath forests along the alpine River Isar using amplified fragment length polymorphisms (AFLP).
• Genetic diversity was, as assumed, slightly higher in gravel bank than in heath forest populations, but genetic differentiation was, in contrast to our expectations, comparable among populations in both habitat types.
• Our study provides evidence for increased genetic diversity under conditions of higher ecological disturbance and increased seed dispersal on gravel banks. Similar levels of genetic differentiation among populations in both habitat types can be attributed to the species' long lifetime, a permanent soil seed bank and gene flow by pollinators among different habitats/locations.

     

Better desiccated than eaten by fish: Distribution of anurans among habitats with different risks to offspring

1. Freshwater species with complex life cycles face a trade-off between the risks of offspring mortality due to desiccation in temporary habitats and due to predators common in long-duration habitats, especially fish. In real-world conditions, intermediate-gradient areas that are highly suitable for some ecological specialists are often limited. I examined the relative significance of drivers of the permanence–predation gradient in habitat selection by pond-breeding anurans.
2. Anuran oviposition was investigated at the community level in fish pond landscapes with only three types of habitat, permanent fish-containing ponds and highly desiccation-prone pools, either fishless or recently emptied of fish and with fish odour (fish cue). Altogether, 65 ponds/pools, interspersed in four clusters, were examined for the presence of egg masses.
3. Bufo bufo was the only species preferring permanent ponds. Egg masses of Rana sp. (Rana arvalis/Rana temporaria) occurred in all fishless pools, and less frequently in the ponds. Bufotes viridis and long larval period species Pelobates fuscus and Hyla orientalis oviposited almost exclusively in fishless pools. Fish-cue pools were avoided by all species. Anuran richness was higher in fishless pools than in permanent ponds. Species distribution between the two habitats was nested, with both common and rare species occurring in fishless pools.
4. The results indicate the primacy of offspring predation risk over hydroperiod constraints in oviposition decision-making by fish-intolerant anurans. The absence of some species from ponds and pools with fish or fish cues shows that non-consumptive interactions may better explain the scarcity of anuran larvae in waters dominated by fish than actual consumption. The strict avoidance of fish habitats despite their proximity to fishless patches indicates fine-scale assessment of predation risk. However, rigid habitat selection against fish predation on offspring may prove maladaptive if habitats with high desiccation risk are the main alternative and are indiscriminately preferred. Maintenance of fishless wetlands with a hydroperiod sufficient to allow completion of metamorphosis should be a conservation priority for anuran diversity in areas where fishless habitats are limited.

     

Colonization of the Mediterranean basin by the vector biting midge species Culicoides imicola: an old story

Understanding the demographic history and genetic make‐up of colonizing species is critical for inferring population sources and colonization routes. This is of main interest for designing accurate control measures in areas newly colonized by vector species of economically important pathogens. The biting midge Culicoides imicola is a major vector of orbiviruses to livestock. Historically, the distribution of this species was limited to the Afrotropical region. Entomological surveys first revealed the presence of C. imicola in the south of the Mediterranean basin by the 1970s. Following recurrent reports of massive bluetongue outbreaks since the 1990s, the presence of the species was confirmed in northern areas. In this study, we addressed the chronology and processes of C. imicola colonization in the Mediterranean basin. We characterized the genetic structure of its populations across Mediterranean and African regions using both mitochondrial and nuclear markers, and combined phylogeographical analyses with population genetics and approximate Bayesian computation. We found a west/east genetic differentiation between populations, occurring both within Africa and within the Mediterranean basin. We demonstrated that three of these groups had experienced demographic expansions in the Pleistocene, probably because of climate changes during this period. Finally, we showed that C. imicola could have colonized the Mediterranean basin in the Late Pleistocene or Early Holocene through a single event of introduction; however, we cannot exclude the hypothesis involving two routes of colonization. Thus, the recent bluetongue outbreaks are not linked to C. imicola colonization event, but rather to biological changes in the vector or the virus.     

Contribution à l'étude des premiers états des Chironomides du Tchad

Résumé Dans cette note, les formes larvaires et nymphales de deux espèces de Chironomides, Tanytarsus nigrocinctus et Chironomus pulcher, sont décrites afin de dégager les caractères systématiques qui permettront ultérieurement l'établissement de clefs de détermination pour les premiers états des Chironomides du Tchad.

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Summary In this note, larves and nymphs of two species of Chironomids, Tanytarsus nigrocinctus and Chironomus pulcher, are described; systematic characters are pointed out permitting in the future confection of key for the dertermination of the first stades of chadian Chironomids.

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Zusammenfassung In dieser Arbeit werden die Larve und Puppe von zwei Chironomiden Arten, Tanytarsus nigrocinctus and Chironomus pulcher, beschrieben. Die artdiagnostischen Merkmale werden hervorgehoben; sie sollen später erlauben, einen dichotomen Schlüssel für die Jugendstadien der Chironomden des Tschadbeckens aufzustellen.

     

Modelling bioenergetic and population-level impacts of invasive bigheaded carps (Hypophthalmichthys spp.) on native paddlefish (Polyodon spathula) in backwaters of the lower Mississippi River

1. While invasions of large rivers by exotic fish species are well documented, assessing actual or potential impacts on native species is a challenge. Rapid assessments may be possible through the application of a combination of bioenergetic and population dynamic models.
2. Paddlefish (Polyodon spathula) is a native species in the central USA with a history of population decline due to waterway development and overharvesting for roe. It is not known whether paddlefish are impacted by resource competition from invasive bigheaded carp populations, including silver (Hypophthalmichthys molitrix) and bighead carp (Hypophthalmichthys nobilis), which have expanded dramatically in the Mississippi River.
3. We used bioenergetic models to project the potential impact of invasive silver and bighead carp on zooplankton density and paddlefish somatic growth in backwater habitat. Bioenergetic outputs were translated to impacts on fecundity, becoming inputs for 50-year metapopulation simulations of backwater habitat connected to the main-stem Mississippi River by episodic flood events.
4. Competition with carp reduced growth and increased the risk of population decline for paddlefish. Impacts increased disproportionately with increased carp abundance and were further exacerbated in scenarios with increased diet overlap or decreased zooplankton abundance.
5. We also analysed paddlefish condition data collected at sites near the lower Mississippi River with varying histories of carp invasion. These data give credence to the bioenergetic model output; paddlefish had reduced body condition at sites with long-established, high-density carp populations.
6. We conclude that invasive bigheaded carps have great potential to reduce paddlefish growth, fecundity, and abundance. The pairing of bioenergetics and population models is likely to be broadly useful in assessing the risks posed by other invasive species.

     

The Mediterranean monk seal Monachus monachus: status,biology, threats,and conservation priorities

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Linking the spatio‐temporal distribution of an edaphic crane fly to its heterogeneous soil environment

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Embryo ecology: Developmental synchrony and asynchrony in the embryonic development of wild annual fish populations

Embryo–environment interactions are of paramount importance during the development of all organisms, and impacts during this period can echo far into later stages of ontogeny. African annual fish of the genus Nothobranchius live in temporary pools and their eggs survive the dry season in the dry bottom substrate of the pools by entering a facultative developmental arrest termed diapause. Uniquely among animals, the embryos (encased in eggs) may enter diapause at three different developmental stages. Such a system allows for the potential to employ different regulation mechanisms for each diapause. We sampled multiple Nothobranchius embryo banks across the progressing season, species, and populations. We present important baseline field data and examine the role of environmental regulation in the embryonic development of this unique system. We describe the course of embryo development in the wild and find it to be very different from the typical development under laboratory conditions. Development across the embryo banks was synchronized within and across the sampled populations with all embryos entering diapause I during the rainy season and diapause II during the dry season. Asynchrony occurred at transient phases of the habitat, during the process of habitat desiccation, and at the end of the dry season. Our findings reveal the significance of environmental conditions in the serial character of the annual fish diapauses.