Relating larval distributions to patterns of oviposition: evidence from lotic hydrobiosid caddisflies

1. To assess the influence of oviposition patterns on distributions of hydrobiosid caddisfly larvae, abundances of three hydrobiosid caddisfly species were estimated in whole reaches with and without suitable oviposition substrata along an upland temperate Australian stream. In addition, within‐reach spatial patterns were examined in relation to known oviposition locations as well as flow characteristics that corresponded to oviposition preferences. 2. Larval abundances in all samples were low relative to previous estimates of egg inputs into reaches. The presence of suitable oviposition substrata at a reach did not influence the abundances of larvae. For one species, benthic samples taken proximate to oviposition substrata revealed a sharp decline in abundance between first and later instars. Larvae of two taxa exhibited flow‐specific habitat preferences mirroring those described as important as cues for oviposition site selection. Previous estimates of egg mass aggregation were also reflected in similarly high levels of larval clumping; however, larval aggregation did not differ between reaches with and without oviposition sites. 3. Collectively, our results suggest that a large difference in the supply of potential recruits does not translate into marked differences in larval abundances of hydrobiosids at the reach level but may account for some variation in larval distribution within a reach and between instars. This evidence is consistent with the notion that (i) posthatching dispersal between reaches is substantial and (ii) mortality of larvae, particularly early instars, is high and (iii) within reach habitat preferences change with larval growth. 4. If adults are unable to lay eggs at reaches without suitable oviposition substrata, then reaches with oviposition substrata may be crucial as a source of recruits elsewhere. Furthermore, high mortality and/or dispersal among first instars signal this as an important part of the life history. Further data on the mortality and dispersal rates of newly hatched larvae would greatly benefit our understanding of the importance of local births in structuring patterns of abundance in stream invertebrates.     

Anomalies in hydropsychid capture nets from polluted streams

• 1 Members of the genus Hydropsyche, net-spinning caddisfly larvae, exhibit anomalies in the structure of their capture nets when collected from stream localities receiving heavy metal and toxic chemical wastes.
• 2 The number of nets showing anomalies was positively correlated with the number of anomalies per net suggesting a graded response to the toxic wastes.
• 3 Frequency of anomalies in a hydropsychid population increased the closer the nets were collected to a pollution source. Anomaly frequency was negatively correlated to total macroinvertebrate community structure as measured by the Shannon function. It is suggested that net anomaly frequeney by hydropsychids may be a useful tool in ecological monitoring.

     

Seston capture by Hydropsyche siltalai and the accuracy of capture efficiency estimates

1. Suspension feeding by caseless caddisfly larvae (Trichoptera) constitutes a major pathway for energy flow, and strongly influences productivity, in streams and rivers. 2. Consideration of the impact of these animals on lotic ecosystems has been strongly influenced by a single study investigating the efficiency of particle capture of nets built by one species of hydropsychid caddisfly. 3. Using water sampling techniques at appropriate spatial scales, and taking greater consideration of local hydrodynamics than previously, we examined the size‐frequency distribution of particles captured by the nets of Hydropsyche siltalai. Our results confirm that capture nets are selective in terms of particle size, and in addition suggest that this selectivity is for particles likely to provide the most energy. 4. By incorporating estimates of flow diversion around the nets of caseless caddisfly larvae, we show that capture efficiency (CE) is considerably higher than previously estimated, and conclude that more consideration of local hydrodynamics is needed to evaluate the efficiency of particle capture. 5. We use our results to postulate a mechanistic explanation for a recent example of interspecific facilitation, whereby a reduction of near‐bed velocities seen in single species monocultures leads to increased capture rates and local depletion of seston within the region of reduced velocity.     

Effects of structural habitat on drift distance and benthic settlement of the caddisfly,Ceratopsyche sparna

We conducted two experiments in flow-through, artificial streams to examine how habitat structure affected drift and benthic resettlement of larval hydropsychid caddisflies (Ceratopsyche sparna). In the first experiment, we quantified drift distance and the number of times larvae re-entered the drift in 9.0 × 0.51-m channels with contiguous patches (ea. 2.5-m long) of biofilm-covered gravel, biofilm-covered cobbles (– Cladophora), and Cladophora-covered cobbles (+ Cladophora). In the second experiment, we tracked nocturnal movements of larvae after benthic settlement in 2.8 × 0.1-m channels, each containing one of the three habitat types. In experiment 1, drift distance was (1) greatest in gravel and lowest in cobbles + Cladophora, (2) inversely related to hydraulic roughness of habitats, (3) independent of body size, and (4) similar for live and dead larvae. Average drift distance was relatively short (<2.5 m), regardless of habitat type. Number of drift re-entries also varied among habitats, being greatest in gravel and lowest in cobbles + Cladophora. No larvae re-entered the drift after settling in Cladophora patches. Results from experiment 2 revealed that drift propensities were higher for larvae in biofilm-covered gravel and cobbles than in cobbles + Cladophora. Larvae remaining in substrate patches (i.e. not drifting) laid fewer draglines in biofilm-covered stones than in Cladophora patches. Extent of benthic movement (i.e., crawling) by non-drifting larvae did not differ significantly among habitats. However, distance moved did differ with flow direction, being 4× greater downstream than upstream. These results highlight how local substrate and hydraulic conditions interact to affect small-scale movements of caddisfly larvae.     

Larval settlement in benthic environments: the effects of velocity and bed element geometry

1. Fluid‐mediated transport can play a key role in determining patterns of distribution and abundance for many benthic invertebrates. One critical challenge in understanding this process is to determine how flow patterns affect larval settlement, especially in those benthic environments where near‐bed flows interact with irregular bed topography to create complex variations in habitat suitability and settlement probability. 2. Boundary‐layer separation over topographical projections on an irregular bed can create two distinct regions of near‐bed flow (i.e., accelerating flow over the forebody and a zone dominated by slower eddies over the aftbody) that may have different effects on larval settlement. 3. We manipulated the flow over a convex roughness element (i.e., hemicylinder) in a flume and examined how the settlement of larvae of the black fly Simulium tribulatum varied with changes in near‐bed velocity and location over the substrate. Larval settlement rate was standardised to correct for variations in larval supply (i.e., among‐trial differences in the concentration of larvae in suspension). 4. Our analyses showed that position on the hemicylinder and near‐bed velocity both affected settlement rate, with a strong interaction effect. In particular, the observed relationship between settlement rate and velocity was negative on the substrate’s forebody and positive on the aftbody. We explore these results by considering potential physical and behavioural mechanisms affecting larval settlement. 5. The presence of a positive relationship between flow and settlement rate in the aftbody may allow settlement on bed elements in habitat where preferred fast‐flow conditions are present, but where settlement would otherwise by hydrodynamically limited. Thus, greater attention to settlement mechanisms in more realistic, topographically complex environments can not only help explain distribution patterns within substrates, but also among substrates and across habitats.     

The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

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Coral larvae settle at a higher frequency on red surfaces

Although chemical cues serve as the primary determinants of larval settlement and metamorphosis, light is also known to influence the behavior and the settlement of coral planulae. For example, Porites astreoides planulae settle preferentially on unconditioned red substrata. In order to test whether this behavior was a response to color and whether other species also demonstrate color preference, settlement choice experiments were conducted with P. astreoides and Acropora palmata. In these experiments, larvae were offered various types of plastic substrata representing three to seven different color choices. Both species consistently settled on red (or red and orange) substrata at a higher frequency than other colors. In one experiment, P. astreoides settled on 100% of red, plastic cable ties but failed to settle on green or white substrata. In a second experiment, 24% of larvae settled on red buttons, more than settled on six other colors combined. A. palmata settled on 80% of red and of orange cables ties but failed to settle on blue in one experiment and settled on a greater proportion of red acrylic squares than on four other colors or limestone controls in a second experiment. The consistency of the response across a variety of plastic materials suggests the response is related to long-wavelength photosensitivity. Fluorescence and reflectance spectra of experimental substrata demonstrated that the preferred substrata had spectra dominated by wavelengths greater than 550 nm with little or no reflection or emission of shorter wavelengths. These results suggest that some species of coral larvae may use spectral cues for fine-scale habitat selection during settlement. This behavior may be an adaptation to promote settlement in crustose coralline algae (CCA)-dominated habitats facilitating juvenile survival.     

Effects of stone topography on abundance of net-building caddisfly larvae and arthropod diversity in an upland stream

We investigated whether small pits on the surfaces of stream stones affected abundances of net-spinning, hydropsychid larvae (Trichoptera) and species diversity in the Acheron River, southeastern Australia. On stream stones, nets occurred more often in small (at least 5 mm width) pits than expected by chance. However, proportionately high numbers of pits were not associated with high abundances of either of two species of hydropsychid larvae (both Asmicridea spp). A colonization experiment, using bricks with 0, 4, 12 or 30 small (diam. 5.5 mm) and large (13 mm) pits drilled into the top surfaces, showed that numbers of nets were related to numbers of pits. Small pits were preferred over large ones and the pits, as a group, were occupied almost entirely by late instars of both species. However, only Asmicridea sp. 1 was more abundant on pitted bricks; Asmicridea sp. 2 showed no response. The differences between the species, and between the stone and brick results, can be explained by different proportions of late instars in the samples. Many I instars probably did not make nets. Hence, higher numbers of larvae on pitted surfaces is expected only when late instars are relatively frequent, as was true for Asmicridea sp. 1 on bricks. There were no effects of pits per se or increased numbers of hydropsychid nets on species diversity or the abundances of other common species. These results contrast with those of other studies, which found stone topography or hydropsychid nets to increase species diversity on stones.     

Coexistence in a guild of filter feeding caddis larvae: do different instars act as different species?

Summary Guild structure and coexistence of filter feeding caddis larvae was studied in three lake-outlet streams in northern Finland. Natural stones were used as sampling units, and for each stone measurements of six microhabitat variables were taken. Species and species instar relationships in the multivariate microhabitat space were analyzed by a method based on Principal Components Analysis. Seven species of filtering caddis larvae were found at the study sites, but detailed analysis was restricted to the four most abundant ones (Polycentropus flavomaculatus, Hydropsyche angustipennis, H. pellucidula, H. saxonica). P. flavomaculatus was the microhabitat generalist of the guild, whereas all the hydropsychids were narrowly specialized in at least one larval instar. In all species, different instars utilized the microhabitat somewhat differentially. Moss content and Froude number were significantly different between instar microhabitats in all comparisons. Ontogenetic niche changes not only separated species instars from each other, but also profoundly changed the biotic as well as abiotic environment of the species at different phases of larval development. This may have important effects on the dynamics of interacting species and, consequently, on the guild structure. While some of the more classical models of coexistence may also be relevant for understanding coexistence in this guild of sessile stream insects, ontogenetic niche changes clearly need to be taken into consideration in future studies of guild structure and species interactions of filter feeding caddisfly larvae.     

Specificity of larval settlement of the Caribbean Orange Icing Sponge,Mycale laevis

The Caribbean sponge Mycale laevis is often found growing in close proximity to living scleractinian corals. This commonly observed sponge–coral association has been considered a mutualism, with the coral providing substratum for the sponge, and the sponge protecting the coral skeleton from boring organisms. We examined the specificity of sponge recruitment to live corals, expecting a positive and specific settlement response if a mutualism exists. Benthic surveys conducted off Key Largo, Florida, and Bocas del Toro, Panama, revealed that individuals of M. laevis grew on substrata that included dead coral and other species of sponges. Selectivity analysis indicated that at three of the four survey sites, M. laevis was not randomly distributed, but associated with live corals more frequently than expected from proportional coral cover. However, settlement assays demonstrated that larvae of M. laevis did not preferentially respond to the presence of live coral. We have previously demonstrated that adults of M. laevis are chemically undefended and readily eaten by spongivorous fishes unless protected by adjacent substrata such as live corals. In overfished areas, where spongivore density is low, the sponge is not selectively distributed near corals. Initial results of settlement experiments with different substrata suggested that larvae of M. laevis responded positively to the presence of the chemically defended sponge Amphimedon compressa, perhaps indicating an associational defense. Further experiments revealed that larvae were reacting to artificially high concentrations of exudates from cut surfaces of Am. compressa; settlement was not enhanced in response to healed pieces of Am. compressa. In addition, the larvae of M. laevis did not selectively respond to live coral or to chemically defended heterospecifics. These results indicate that the commonly observed proximity of M. laevis to live corals is not driven by larval settlement behavior, but instead by post‐settlement mortality due to predation.     

Onshore–offshore variations of copepod community in northern South China Sea

Previous studies which have tested the feeding preferences of shredders for fungal species and the food quality of fungi used detritus uniformly colonized by a fungus, which is not the case for decaying leaves in streams. It is not known whether shredders in different development stages exhibit variations in feeding preference and larval performance. This study examined the feeding preferences and the growth of the third and the fifth instars of Pycnopsyche gentilis larvae using fungal-colonized patches and whole leaves, respectively, having different fungal species compositions (Alatospora acuminata, Anguillospora filiformis, Articulospora tetracladia, Tetrachaetum elegans, and all species combined). The aquatic hyphomycetes used were co-dominant on leaves in the stream inhabited by the caddisfly. During 14 d of feeding, the larvae of both instars did not show significant differences in feeding preferences for the patches growing on oak leaves, although the third instar larvae were slightly more selective than the fifth instar larvae. When fed with maple leaves for 18 d, larval growth rates, gross growth efficiencies, and survivorship were not significantly different among the fungal treatments. However, the larval growth of both instars fed with fungal-colonized leaves was always significantly greater than the growth of larvae fed with diets of uncolonized leaves. The third instar larvae grew faster than the fifth instar larvae, but the growth efficiencies of the two instars were similar. These results suggest that P. gentilis larvae exhibit less selectivity in their feeding than other caddisfly shredders that have been examined and that the dominant fungi colonizing leaves in their habitat are similar in palatability and food quality for this shredder. Handling editor: B. Oertli     

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.

     

The influence of moss on grazers in high‐altitude streams: food,refuge or both?

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Resilience of aquatic net‐spinning caddisfly silk structures to common global stressors

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Disturbance and investment: developmental responses of tropical lotic midges to repeated tube destruction in the juvenile stages

Abstract. 1. Investment in silken tubes may give tubicolous chironomid larvae more to lose than non‐tube building taxa when physical disturbances or the actions of other organisms induce dispersal. In this study, two chironomid species from flood‐prone Australian tropical streams were subjected to repeated experimental destruction of their larval and pupal tubes in laboratory experiments, with life‐history parameter responses recorded. 2. When subjected to the tube destruction treatments, Echinocladius martini (Orthocladiinae), whose larvae construct branched loose silken tubes in fast flow, showed no reduction in survivorship to adult, whereas Polypedilum australotropicus (Chironominae), which builds compact tightly woven tubes incorporating detrital material within pool leaf packs, showed reduced survivorship of about 25%. 3. For both species, tube destruction reduced pupal duration, regardless of whether destruction occurred in the larval stage only, or in both the larval and pupal stages. This may be a response to the risks of exposure outside tubes in nature, particularly for pupae, which are incapable of spinning silk. 4. Tube destruction caused little or no reduction in adult size, longevity, or female fecundity (oocyte number) for either species, but oocyte development and size was retarded in E. martini females, reflecting the energetic and nutritive costs of the tube destruction treatments. 5. Contrasts in the responses of the two species reflect differences in tube‐building behaviour and in situ habitat use. These results demonstrate that tube loss can affect the fitness of individual chironomids, though both species proved generally resilient, which may reflect their long‐term exposure to variable tropical lotic conditions.     

The influence of resident adults on larval settlement: experiments with four species of ascidians

Residents within any community can affect the larval settlement of both their own and other species. In marine sessile communities resident adults can affect larval settlement by preying on settling larvae, removing or adding space for the larvae to colonize, or stimulating or prohibiting larval settlement on available substratum nearby. To examine those processes by which residents affect settlement, we exposed experimental substrata with three densities of adults of a single species at a site in eastern Long Island Sound, USA for a 24-h period. Four species of common ascidians, Botryllus schlosseri (Pallas), Botrylloides diegensis Ritter and Forsyth, Diplosoma macdonaldi Herdman, and Molgula manhattensis (De Kay), were used in 11 separate experiments. Few individuals of any species settling attached to the surfaces of these species and this resulted in the main effect of these residents being the usurpation of space and the restricting of settlement to unoccupied areas. A model is also presented to explain the apparent aggregated settlement of several species in open areas adjacent to the resident ascidians. From this model we suggest that the aggregated settlement can result from limited larval mobility such that some larvae that contact and reject the resident species as settlement sites may subsequently contact open surfaces of the same substratum and increase settlement densities there over those observed on control substrata. Finally, settlement data for several species indicate that Molgula may influence settlement by preying on larvae.     

Habitat differentiation in the early life stages of simultaneously mass-spawning corals

The settlement process of coral larvae following simultaneous mass-spawning remains poorly understood, particularly in terms of population and community parameters. Here, the larval settlement patterns of Acropora corals, which are the most diverse genera of scleractinian corals at the species (haplotype) level, were investigated within a single subtropical reef. Across a 4-year period (2007–2010), the mitochondrial and nuclear molecular markers of 1,073 larval settlers were analyzed. Of the 11 dominant haplotypes of recruited populations, nine exhibited non-random patterns of settlement distribution. This result suggests that the actual habitat segregation starts during the early swimming larval stages of their life history, rather than by natural selection after random settlement. In addition, the presence of a depth-related settlement pattern supports that species-specific vertical zonation of coral larvae may play a role in the establishment of habitat segregation. Moreover, in some species that showed a preference toward the shoreward area of the bay, the settlement pattern was consistent with that of the adult distribution. This result indicates that the gametes were not mixed between fore and back reefs in the period from fertilization to settlement during the mass-spawning event, even within a single small reef. Another compatible hypothesis of this pattern is that the larvae are able to recognize various types of environmental information, facilitating the selection of optimal micro-habitats. Overall, Acropora coral larvae that are produced from a simultaneous mass-spawning event may have adapted to complex reef topography by means of multi-step habitat selection at settlement, corresponding to different spatial scales.     

Non-Additive Increases in Sediment Stability Are Generated by Macroinvertebrate Species Interactions in Laboratory Streams

Previous studies have shown that biological structures such as plant roots can have large impacts on landscape morphodynamics, and that physical models that do not incorporate biology can generate qualitatively incorrect predictions of sediment transport. However, work to date has focused almost entirely on the impacts of single, usually dominant, species. Here we ask whether multiple, coexisting species of hydropsychid caddisfly larvae have different impacts on sediment mobility compared to single-species systems due to competitive interactions and niche differences. We manipulated the presence of two common species of net-spinning caddisfly (Ceratopsyche oslari, Arctopsyche californica) in laboratory mesocosms and measured how their silk filtration nets influence the critical shear stress required to initiate sediment grain motion when they were in monoculture versus polyculture. We found that critical shear stress increases non-additively in polycultures where species were allowed to interact. Critical shear stress was 26% higher in multi-species assemblages compared to the average single-species monoculture, and 21% greater than levels of stability achieved by the species having the largest impact on sediment motion in monoculture. Supplementary behavioral experiments suggest the non-additive increase in critical shear stress may have occurred as competition among species led to shifts in the spatial distribution of the two populations and complementary habitat use. To explore the implications of these results for field conditions, we used results from the laboratory study to parameterize a common model of sediment transport. We then used this model to estimate potential bed movement in a natural stream for which we had measurements of channel geometry, grain size, and daily discharge. Although this extrapolation is speculative, it illustrates that multi-species impacts could be sufficiently large to reduce bedload sediment flux over annual time scales in streams where multiple species of caddisfly are present.     

Maternal behaviours may explain riffle‐scale variations in some stream insect populations

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