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.     

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.     

A new anhydrobiotic midge from Malawi,Polypedilum pembai sp.n. (Diptera: Chironomidae), closely related to the desiccation tolerant midge,Polypedilum vanderplanki Hinton

The sleeping chironomid (Polypedilum vanderplanki Hinton) lives on temporary rock pools in the semi‐arid tropical regions of Africa. Its larvae are able to survive the dry season in a completely desiccated ametabolic state known as anhydrobiosis. So far, P. vanderplanki was the only species among all insects showing demonstrated anhydrobiotic ability. Here, we show that a new related species originating from Malawi, Polypedilum pembai sp.n. , is also anhydrobiotic and that its desiccation tolerance mechanism is probably similar to what is observed in P. vanderplanki. The new species, P. pembai sp.n. , is described with special attention to the common and different morphological features, compared with P. vanderplanki. Phylogenetic analysis showed that both species are closely related, suggesting that anhydrobiosis evolved only once in their common ancestor about 49 Ma somewhere in Africa, before the divergence of two species, one in the sub‐Saharan area and another in southeastern Africa.     

Density and distribution in laboratory populations of midge larvae (Chironomidae: diptera)

The troglobitic amphipod crustacean Crangonyx antennatus occupies mud-bottom pools and small, gravel-bottom streams in caves in the southern Appalachians. One large, mud-bottom pool population in Lee Co., Virginia was observed periodically from 1967 to 1975. Amphipods in this population frequently burrowed into the soft mud substrate, where they were able to survive desiccation during periods when the pool dried up. Animals kept in the laboratory also burrowed and survived desiccation during an experiment which simulated drought conditions similar to those observed in caves. Amphipods collected from both pool and stream habitats burrowed, thereby indicating that stream-adapted populations of this species still retain sufficient flexibility to survive under variable environmental conditions that might be encountered in nature. It is concluded that burrowing provides a means of survival for C. antennatus when it is sometimes exposed to drought conditions in cave pool habitats and also provides protection from potential terrestrial predators under similar conditions. Burrowing also offers amphipods protection from aquatic predators (such as salamander larvae) during normal water levels and possibly allows juveniles a means of escaping cannibalism by adults.     

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.     

Long-term movements of self-marked caddisfly larvae (Trichoptera: Sericostomatidae) in a California coastal mountain stream

1. Over 140 000 larvae of the case-building caddisfly Gumaga nigricula were self-marked as they incorporated glitter into small portions of their cases while reared in streamside troughs. These marked individuals were released into stream pools and their movements monitored in the dry season, when base flow was low and no spates occurred, and in the wet season when base flow was high and several spates occurred. 2. Of the 9,000–10 000 larvae released in each of two stream pools in the dry season, 4–20% (i.e. 377–1817 marked individuals) were observed on three sampling dates (4, 11 and 24 d after release). Most larvae (87–93%) remained within 4 m up- or downstream of the release line after 24 d. No larvae were found outside of the release pools, even after 37 d. 3. Of the > 120 000 larvae released in one stream pool near the beginning of the wet season, 408 larvae were recaptured 130–167 d later, a period that included 30 days of high flow associated with six spates. Estimated survivorship over this period was 0.7–6.2%; there was no relationship between survival and larval size at release. Most (75%) recaptured larvae were found in the pool where they were originally released. The remaining larvae were found downstream of the release pool. Larvae had generally dispersed only a short distance downstream of the release pool (median = 18 m, maximum = 222 m). In addition, four marked pupae were later found 436 m downstream of the release pool. 4. These results illustrate the sedentary nature of larval G. nigricula as well as the important role that high flow events play in larval mortality and dispersal. These case-building larvae move very little during low flow periods, even when food resources appear limiting. In contrast, the frequency and distance of larval dispersal are much greater during periods with high flow. 5. Our observations for G. nigricula support previously published inferences that larval dispersal within a stream can be limited for some aquatic insects. However, our observations also suggest that, even for a relatively sedentary species like G. nigricula, larval dispersal during periods with high flow may contribute significantly to gene flow within a stream reach.     

Survival of caddis larvae in relation to their case material in a group of temporary and permanent pools

1. The occurrence of larvae of Limnephilus coenosus and L. vittatus (Trichoptera) was recorded in 103 rock pools both before and after the dry periods in spring–summer (April–May and July–early August), when a great number of the pools dried out. In early spring, 84% of the pools sampled contained larvae of at least one of the species. Limnephilus coenosus larvae were present in a higher proportion of pools that later dried out than in the permanent ones, while L. vittatus larvae were predominantly found in permanent pools. Larvae of both species were found together in 31% of the inhabited pools.
2. Laboratory experiments, designed to elucidate differential survival under drought conditions, demonstrated that the organic cases of L. coenosus larvae held water more efficiently than did the mineral L. vittatus cases. Full-grown (fifth instar) L. coenosus larvae also survived dry conditions better than did L. vittatus larvae. The higher survival of L. coenosus was due to a combination of drought-resistant case material and a higher survival ability of the larvae themselves.
3. Limnephilus vittatus larvae were restricted by their low capability to withstand desiccation, although case material and food were available for L. vittatus larvae in all pools. Absence of case material and high-quality food in many pools, however, restricted the presence of L. coenosus larvae, but this species did not suffer from heavy mortality because of desiccation and all pool populations of L. coenosus survived until pupation.
4. The differences in species composition in these temporary and permanent pools can be explained by the differential site selection by ovipositing females, as well as by larval survival. The intrinsically greater survival of L. coenosus larvae during drought, together with the water-retaining properties of the cases, allowed this species to exploit stressful and risky habitats, such as temporary pools.     

Survival of caddis larvae in relation to their case material in a group of temporary and permanent pools

1. The occurrence of larvae of Limnephilus coenosus and L. vittatus (Trichoptera) was recorded in 103 rock pools both before and after the dry periods in spring–summer (April–May and July–early August), when a great number of the pools dried out. In early spring, 84% of the pools sampled contained larvae of at least one of the species. Limnephilus coenosus larvae were present in a higher proportion of pools that later dried out than in the permanent ones, while L. vittatus larvae were predominantly found in permanent pools. Larvae of both species were found together in 31% of the inhabited pools.
2. Laboratory experiments, designed to elucidate differential survival under drought conditions, demonstrated that the organic cases of L. coenosus larvae held water more efficiently than did the mineral L. vittatus cases. Full-grown (fifth instar) L. coenosus larvae also survived dry conditions better than did L. vittatus larvae. The higher survival of L. coenosus was due to a combination of drought-resistant case material and a higher survival ability of the larvae themselves.
3. Limnephilus vittatus larvae were restricted by their low capability to withstand desiccation, although case material and food were available for L. vittatus larvae in all pools. Absence of case material and high-quality food in many pools, however, restricted the presence of L. coenosus larvae, but this species did not suffer from heavy mortality because of desiccation and all pool populations of L. coenosus survived until pupation.
4. The differences in species composition in these temporary and permanent pools can be explained by the differential site selection by ovipositing females, as well as by larval survival. The intrinsically greater survival of L. coenosus larvae during drought, together with the water-retaining properties of the cases, allowed this species to exploit stressful and risky habitats, such as temporary pools.     

What to do if streams go dry? Behaviour of Balkan Goldenring (Cordulegaster heros,Odonata) larvae in a simulated drought experiment in SW Hungary

1. In case of dryings, the hyporheic zone is one of the most important refugia for stream macroinvertebrate communities, including the few Odonata species living in these habitats, such as Cordulegaster species. There is no information on the desiccation resistance strategies and methods of any members of the genus, including Cordulegaster heros.
2. We hypothesised that the larvae use burrowing behaviour to survive droughts. In this study, beyond recording the survival rates of the larvae, we tested the effects of the sediment particle size and the body size of the larvae on burrowing behaviour in a 3-week-long simulated drought experiment in an indoor artificial stream system.
3. Eighty larvae were involved in the experiment, from which 60 were treated with drought, and 20 served as controls. Larvae were put into flowing water, into separate special compartments; 1 day later, the flow was ceased, and then, the water level was gradually decreased for 3 weeks.
4. Approximately 15% of larvae could survive the 3 weeks of drying. The survival probability of drought-treated larvae was significantly increased if animals burrowed into the sediment. In addition, the survival probability was higher in case of fine substrate material. Size of the larvae only affected the depth of the burrowing, not the survival rate.
5. However, two-thirds of the larvae did not dig into the sediment, which implies that surviving via burrowing is not the only mechanism of the species to withstand dry periods.

     

Refugia and habitat partitioning among midges (Diptera: Chironomidae) in rain-pools

ABSTRACT.

• 1 Rain-pools on granite exposures are common in tropical Africa and they are inhabited by the larvae of two midge species, Chironomus imicola Kieffer and Chironomus pulcher Wiedemann.
• 2 These pools differ from other ephemeral island habitats such as carrion, fallen fruit, sap flows, mushrooms and dung in their high degree of spatial predictability. Most pools are predictably inhabited by either C.imicola or C.pulcher. This finding is considered in the light of contemporary competition theory.
• 3 An arrangement whereby the same species repeatedly invades the same temporary habitat indicates that there is something about the habitat itself causing the discrimination. I suggest that degree of exposure to sunlight may provide a cue allowing females to discriminate between pools as oviposition sites.
• 4 Both species survive the long dry season by retreating to pools of river water left by receding rivers. Dispersal to rain-pools by emerging adults occurs during the following season with imicola occupying rain-pools remote from shaded rivers. Appropriately, measured against pulcher, the biology of imicola is that of a colonizing species.
• 5 The possibility is discussed that the chironomids in rock-pools do not survive to breed. Instead, these populations may be maintained externally, for example by invasions from breeding populations in the Great Lakes of Africa, where larval densities may be too low to be detected by sampling. The evolutionary implications of such a situation are considered.

     

Dried-up zone as a temporal stock of chironomid larvae: survival periods and density in a reservoir bank

The survival period, community composition and density of chironomid larvae were investigated at the reservoir bank in Yachiyoko Lake (Haji Dam), Hiroshima, Japan to verify their survivability in a dried-up zone. We define “dried-up zone” as the one that had been exposed to drought due to drawdown by evaporation and/or drainage. The larvae were collected from sediments at different altitudes along a transectional line. After larval rearing, 189 indiv. m⁻² of adults emerged even from the sediment of a 7-day drought (17.1% sedimental water content). The proportion of Tanytarsus oyamai Sasa in the community tended to be larger with dry periods, indicating that this species has a higher drought tolerance than other species. The mean density in the dried-up zone was estimated as 886.4 indiv. m⁻². Biomass is generally estimated by using the samples of only underwater sediment. These results show that some species have the potential to survive for at least one week in a dried-up zone and suggest that the biomass is likely to be highly underestimated for the entire ecosystem. A pathway of matter flow from aquatic to terrestrial ecosystems via the larvae in a dried-up zone should be taken into consideration.     

Occurrence of additional Zoea-VI larvae in the mud crab, Scylla paramamosain (Estampador), reared in the laboratory

Mud crabs, Scylla spp. , are commercially important in many Indo-Pacific countries. The larval development of mud crabs has been reported previously as five zoeal and one megalopal stages. This paper reports larval rearing experiments that revealed variability in larval developmental stages in the mud crab Scylla paramamosain, one of four mud crab species. In addition to normal five zoeal stages, an alternative pathway of developing through six zoeal stages was observed for the crab. There were evidences suggested that the appearance of the additional Zoea-VI larvae was associated with unfavourable dietary conditions, including poor quality of diet, inadequate quantity of dietary supply and a period of starvation for newly hatched larvae. Based on exuviae and larval specimens, the morphology of the additional Zoea-VI larvae was described.     

Factors Inducing Successful Anhydrobiosis in the African Chironomid Polypedilum vanderplanki: Significance of the Larval Tubular Nest

The African chironomid Polypedilum vanderplanki exhibits anhydrobiosis,i.e., the larvae can survive complete desiccation. Recoveryrate and trehalose content were investigated in larvae desiccatedslowly or at a rate more than 3 times faster. Upon slow desiccation(evaporation rate 0.22 ml day^–1) larvae synthesized 38µg trehalose/individual before complete desiccation, andall of them recovered after rehydration, whereas larvae thatwere dehydrated quickly (evaporation rate 0.75 ml day^–1)accumulated only 6.8 µg trehalose/individual and noneof them revived after rehydration. In the pools that are theirnatural habitat P. vanderplanki larvae make tubes by incorporatingdetritus or soil with their sticky saliva. This tubular structureis a physical barrier not only to protect the larva from naturalenemies but also induces successful anhydrobiosis by reducingthe dehydration rate. When larvae were dehydrated with 100 µldistilled water (DW) in soil tubes, they accumulated 37 µgtrehalose/individual and more than half of them could reviveafter rehydration, whereas larvae without tubes accumulatedlower level of trehalose and none recovered after rehydration.     

Unexpected collective larval dispersal but little support for sweepstakes reproductive success in the highly dispersive brooding mollusc Crepidula fornicata

In many marine invertebrates, long‐distance dispersal is achieved during an extended pelagic larval phase. Although such dispersal should result in high gene flow over broad spatial scales, fine‐scale genetic structure has often been reported, a pattern attributed to interfamilial variance in reproductive success and limited homogenization during dispersal. To examine this hypothesis, the genetic diversity of dispersing larvae must be compared with the postdispersal stages, that is benthic recruits and adults. Such data remain, however, scarce due to the difficulty to sample and analyse larvae of minute size. Here, we carried out such an investigation using the marine gastropod Crepidula fornicata. Field sampling of three to four larval pools was conducted over the reproductive season and repeated over 3 years. The genetic composition of larval pools, obtained with 16 microsatellite loci, was compared with that of recruits and adults sampled from the same site and years. In contrast to samples of juveniles and adults, large genetic temporal variations between larval pools produced at different times of the same reproductive season were observed. In addition, full‐ and half‐sibs were detected in early larvae and postdispersal juveniles, pointing to correlated dispersal paths between several pairs of individuals. Inbred larvae were also identified. Such collective larval dispersal was unexpected given the long larval duration of the study species. Our results suggest that each larval pool is produced by a small effective number of reproducers but that, over a reproductive season, the whole larval pool is produced by large numbers of reproducers across space and time.     

Invertebrate community dynamics and insect emergence in response to pool drying in a temporary river

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Priority effects of the early breeding fire salamander on the late breeding banded newt

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The effects of cannibalism on Amazonian poison frog egg and tadpole deposition and survivorship in Heliconia axil pools

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Effects of dehydration rate on physiological responses and survival after rehydration in larvae of the anhydrobiotic chironomid

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