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

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Ecology and population structure of the bayou darter, <Emphasis Type="Italic">Etheostoma rubrum</Emphasis>: disjunct riffle habitats and downstream transport of larvae

The bayou darter, Etheostoma rubrum (Percidae), is endemic to the Bayou Pierre system in Mississippi. Adult and juvenile E. rubrum occupy swift, shallow riffles or runs over coarse gravel and pebble substrata. Habitat requirements of larval and post-larval stages, and the role of downstream dispersal of larvae in colonizing riffles are poorly known. The potential for movement and the high level of habitat specificity for the discontinuous riffle habitat suggest that E. rubrum may comprise a metapopulation of linearly arranged local populations. The greatest population densities of E. rubrum occur in the upper reaches of Bayou Pierre. We hypothesized that metapopulation structure of E. rubrum may include source–sink dynamics, whereby downstream areas are a sink for larvae/early juveniles originating upstream. We tested hypotheses that a transport mechanism, larval drift, occurred in E. rubrum, and that downstream riffles showed characteristics of population sinks. We captured larval stages of E. rubrum in cross-sectional and longitudinal drift nets, and rates of drift tended to increase during the day. Larval E. rubrum (n=19) occurred in samples above and below riffle areas, with no differences among longitudinal drift nets placed above and below riffles. Thus, larval drift is a viable transport mechanism. Relative abundance of adults and juveniles declined from upstream to downstream, and inter-riffle distances increased with cumulative downstream distance. However, both predictions of the source–sink hypothesis were not supported. The distribution of size classes did not change between upstream and downstream riffles nor did the mean size-adjusted body mass.     

Large-scale manipulation of mayfly recruitment affects population size

Recruitment establishes the initial size of populations and may influence subsequent population dynamics. Although strong inference can be made from empirical relationships between recruitment and population sizes, a definitive test of recruitment limitation requires manipulating recruitment at relevant spatial and temporal scales. We manipulated oviposition of the mayfly Baetis bicaudatus in multiple streams and measured the abundance of late-stage larvae at the end of the cohort. Based on fundamental knowledge of mayfly behavior, we increased, eliminated, or left unmodified preferred mayfly oviposition sites in 45-m reaches of streams (N = 4) of one high-altitude drainage basin in western Colorado, USA. We compared egg densities before (2001) and after the manipulation (2002) using paired t tests and compared larval densities before and after the manipulation among treatments using repeated measures analysis of variance. This manipulation altered not only egg densities, but also larval abundances 1 year later. Compared to the previous year, we experimentally increased egg densities at the addition sites by approximately fourfold, reduced egg densities to zero in the subtraction sites, and maintained egg densities in the control sites. After the manipulation, larval densities increased significantly by a factor of approximately 2.0 in the addition sites and decreased by a factor of approximately 2.5 in the subtraction sites. This outcome demonstrates that dramatic changes in recruitment can limit larval population size at the scale of a stream reach, potentially masking previously observed post-recruitment processes explaining the patterns of variation in abundance of a stream insect. Furthermore, our results emphasize the importance of preferred oviposition habitats to population sizes of organisms.     

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.     

Dispersal in drift-prone macroinvertebrates: a case for density-independence

1. Studies of dispersal of macroinvertebrates in streams and rivers tend to be focused on drift, whilst benthic movements are usually considered to be less important.
2. Field-enclosure experiments with the mayfly Baetis rhodani indicate that net dispersal in this species is simply a proportional loss of individuals from the benthos.
3. Neither net upstream or downstream movements exhibited evidence of density-dependence in the form of curvilinear relationships between initial and final densities.
4. The net number of animals moving upstream did not differ significantly from the net number moving downstream.
5. The probable mechanisms behind density-independent dispersal are discussed, as are the implications for our understanding of population dynamics in relation to invertebrate drift.     

Oviposition site selectivity of some stream-dwelling caddisflies

Population dynamics depends upon the spatial distribution of individuals in heterogeneous environments. The various processes surrounding insect oviposition are central to understanding their population dynamics because the choice of oviposition site ultimately influences the survivorship and spatial distribution of their progeny. Aquatic insects are often assumed to have non-selective oviposition habits, but empirical data are scarce and selective oviposition may be quite common. We quantitatively sampled egg masses of stream-dwelling caddisflies (Trichoptera) that specialise in egg-laying on hard substrata underwater, in order to characterise oviposition site selectivity and test for communal oviposition. In a field survey of two Scottish streams, we sampled egg masses of three species, Polycentropus flavomaculatus, Hydropsyche siltalai, Rhyacophila dorsalis, with the aim of testing whether egg mass abundance varied with current (riffles vs. pools), location within the channel (margins vs. centre) and rock exposure (emergent vs. fully submerged). In one stream, we captured adults landing on emergent rocks and assessed whether females were modified morphologically for swimming. The egg masses of two species (P. flavomaculatus, H. siltalai) occurred primarily on submerged rocks in pool margins, and adult females had legs modified for swimming. In contrast, egg masses of R. dorsalis were most abundant on the underside of emergent rocks in riffles, and females were not modified for swimming. Communal oviposition was evident for all three species, with most egg masses aggregated on the minority of potential rocks. How females locate oviposition sites and the consequences of these highly specialised oviposition behaviours to the survival and spatial distribution of larvae now require investigation. The effects of these behaviours on population dynamics are likely to differ from terrestrial herbivores because oviposition sites are not food resources for these aquatic species.     

The influence of leaf age on the oviposition preference of Chrysophtharta bimaculata (Olivier) and the establishment of neonates

1 The degree of discrimination shown by a herbivore when selecting oviposition sites has been suggested as a key factor to understanding herbivore population dynamics. Chrysophtharta bimaculata (Coleoptera: Chrysomelidae) is a primary pest of Tasmanian eucalypt forests and can cause severe defoliation. Previous work suggests that females show discrimination when selecting oviposition sites. Our aim was to test the degree of oviposition discrimination exhibited by C. bimaculata with regards to leaf toughness, a character that is critical to neonate survival. 2 We conducted an experiment examining the leaf toughness critical for neonate survival and found that significant larval mortality occurs above a toughness of 46.9 g. We also determined that the maximum toughness of leaves upon which larvae established in the field was 48.2 g, supporting the laboratory result. 3 Field surveys showed that although the majority of eggs were laid on leaves suitable for larval establishment, many eggs were laid on unsuitable, tougher leaves. However, all eggs were normally placed within 20 cm of suitable leaves and glasshouse trials demonstrated the neonates could move this distance without mortality occurring. 4 We conclude that egg batch distribution and larval performance of C. bimaculata will influence the population dynamics of C. bimaculata in two ways. Firstly, the availability of expanding/newly expanding leaves of eucalypt hosts will determine larval carrying capacity. Secondly, at a more localized level, the deposition of large numbers of egg batches on both suitable and unsuitable leaves followed by successful neonate migration increases the risk of resource depletion and poor larval development.     

Oviposition and larval dispersal of the common armyworm,Mythimna convecta (Walker) (Lepidoptera: Noctuidae)

The common armyworm, Mythimna convecta (Walker), is a pest of cereal crops and pasture grasses in Australia. During autumn, egglaying in grasslands commonly occurs before plant growth commences. The possible association between oviposition and dried grasses was investigated in field studies of larval distribution in a pasture and a crop habitat, and in laboratory studies of oviposition site preferences and the mode of dispersal of newly hatched first instar larvae. A comparison of a green grass pasture with and without a component of dried grass showed that highest densities of M. convecta larvea. were found in the former. In the laboratory, egg batches were laid between plant parts in close contact, such as between the stem and leaf axil and between seeds in the seed head. The crevices in dried plant material and seed-heads were significantly more acceptable for oviposition than in green foliage. Oviposition in dried plant material enables M. convecta to rapidly colonize ephemeral grasslands immediately after rains when the neonate larvae can disperse and feed on new vegetative growth. Shortly after hatching, 93% of first instar larvae used fine silken threads for aerial dispersal in 0.5 m s^-1 air currents and 43% were blown a distance of more than 1.0 m. In the field, sticky traps caught larvae dispersing in all directions from a central source, and indicated that approximately 30% were transported 2 m after 7 days of light to medium breezes. Terminal velocities of newly hatched larvae were 20–130 cm s^-1, depending on the length of the trailing thread.     

Linking movement and oviposition behaviour to spatial population distribution in the tree hole mosquito Ochlerotatus triseriatus

1. Researchers often use the spatial distribution of insect offspring as a measure of adult oviposition preferences, and then make conclusions about the consequences of these preferences for population growth and the relationship between life-history traits (e.g. oviposition preference and offspring performance). However, several processes other than oviposition preference can generate spatial patterns of offspring density (e.g. dispersal limitations, spatially heterogeneous mortality rates). Incorrectly assuming that offspring distributions reflect oviposition preferences may therefore compromise our ability to understand the mechanisms determining population distributions and the relationship between life-history traits. 2. The purpose of this study was to perform an empirical study at the whole-system scale to examine the movement and oviposition behaviours of the eastern tree hole mosquito Ochlerotatus triseriatus (Say) and test the importance of these behaviours in determining population distribution relative to other mechanisms. 3. A mark-release-recapture experiment was performed to distinguish among the following alternative hypotheses that may explain a previously observed aggregated distribution of tree hole mosquito offspring: (H(1)) mosquitoes prefer habitats with particular vegetation characteristics and these preferences determine the distribution of their offspring; (H(2)) mosquitoes distribute their eggs randomly or evenly throughout their environment, but spatial differences in developmental success generate an aggregated pattern of larval density; (H(3)) mosquitoes randomly colonize habitats, but have limited dispersal capability causing them to distribute offspring where founder populations were established; (H(4)) wind or other environmental factors may lead to passive aggregation, or spatial heterogeneity in adult mortality (H(5)), rather than dispersal, generates clumped offspring distributions. 4. Results indicate that the distribution of tree hole mosquito larvae is determined in part by adult habitat selection (H(1)), but do not exclude additional effects from passive aggregation (H(4)), or spatial patterns in adult mortality (H(5)). 5. This research illustrates the importance of studying oviposition behaviour at the population scale to better evaluate its relative importance in determining population distribution and dynamics. Moreover, this study demonstrates the importance of linking behavioural and population dynamics for understanding evolutionary relationships among life-history traits (e.g. preference and offspring performance) and predicting when behaviour will be important in determining population phenomena.     

Hydrologic and behavioral constraints on oviposition of stream insects: implications for adult dispersal

The supply of recruits plays an important role in plant and animal population dynamics, and may be governed by environmental and behavioral constraints on animals. Mated females of the mayfly genus Baetis alight on rocks protruding from streams, crawl under water and deposit a single egg mass under a rock. We surveyed oviposition and emergence of a bivoltine population of B. bicaudatus in multiple stream reaches in one high-altitude watershed in western Colorado over 3 years to establish qualitative patterns at a regional scale (entire watershed), and quantitative patterns over six generations at a local scale (one stream reach). We also measured characteristics of preferred oviposition substrates, performed experiments to test hypotheses about cues used by females to select oviposition sites, and measured mortality of egg masses in the field. Our goals were to determine whether: (1) hydrologic variation necessitated dispersal of females to find suitable oviposition sites; (2) the local supply of females could provide the supply of local recruits; and (3) local recruitment determined the local production of adults. The onset of oviposition corresponded with the decline of spring run-off, which differed dramatically among years and among sites within years. However, eggs appeared before any adults had emerged in 8 of 22 site-years, and adults emerged 2-3 weeks before any eggs were oviposited in 3 site-years. Furthermore, the size distribution of egg masses differed from that predicted by the size distribution of females that emerged from seven of nine stream reaches. Protruding rocks and eggs appeared earlier each summer in smaller tributaries than in larger mainstream reaches, suggesting that hydrologic and behavioral constraints on oviposition may force females to disperse away from their natal reach to oviposit, and possibly explain the predominantly upstream flight of Baetis females reported in other studies. Local oviposition rates in one third-order stream-reach increased rapidly as soon as substrates protruded from the water surface, and females preferred large rocks that became available early in the flight season. However, females oviposited on <10% of all available rocks, and <65% of preferred rocks as determined by an empirical model. These data indicated that the timing of appearance of suitable oviposition sites determined the phenology of local recruitment, but that preferred oviposition sites were not saturated. Thus, the magnitude of local recruitment was not limited by the absolute abundance of preferred oviposition sites. Only 22% of egg masses observed in the field suffered mortality during their embryonic development, and per capita Baetis egg mass mortality was significantly lower on rocks with higher densities of egg masses. Thus, we suspect that specialized oviposition behavior may reduce the probability of egg mortality, potentially compensating for the costs of dispersal necessary to locate suitable oviposition sites. Finally, the number of adults that emerged at one stream reach was independent of the number of egg masses oviposited over six generations of Baetis; and local recruitment was not a function of the number of adults of the previous generation that emerged locally. The patterns of oviposition and emergence of Baetis found in this study are consistent with the following hypotheses. Recruitment of eggs in a stream reach is not limited by the local supply of adults, but is a function of the regional supply of dispersing adults, which are constrained by the spatial and temporal distribution of preferred oviposition habitat. Furthermore, subsequent local production of adults is not a function of the supply of recruits, arguing for post-recruitment control of local populations by processes operating in the larval stage (e.g., predation, competition, dispersal, disturbance). Processes affecting larval and adult stages of Baetis act independently and at different scales, thereby decoupling local population dynamics of successive generations.     

Terrestrial–aquatic transitions: Local abundances and movements of mature female caddisflies are related to oviposition habits but not flight capability

1. Movement behaviours of adult aquatic insects can produce distinct spatial distribution patterns. Studies of adult abundance with distance away from water bodies are common and may invoke flight capability to explain species differences. In contrast, distribution patterns along river channels are poorly described, but are no less important for understanding population dynamics. Longitudinal patterns in adult abundance along short river lengths may differ between sexes and at different life stage transitions between aquatic and terrestrial environments, i.e. at emergence and oviposition. Flight capability is unlikely to influence longitudinal patterns created at emergence, but may influence local abundances of mature females seeking to lay eggs. We tested hypotheses about how local abundances of mature females might differ according to oviposition habits and flight capability.
2. We surveyed abundances of mature female caddisflies at adjacent riffle–pool pairs along short river lengths with homogeneous riparian cover. Our survey included nine species in three families (Hydrobiosidae, Leptoceridae, Hydropsychidae), which encompassed multiple different oviposition habits and a range of wing sizes and shapes. Several of the species oviposit preferentially in riffles. Accordingly, we tested for differences in female abundance between channel units (adjacent riffle–pool pairs). We also tested whether females attained higher abundances in some places along channels than others (i.e. over larger spatial scales and regardless of channel unit) which imply movements along the channel and aggregation in some locations. Wing morphology was used as a proxy measure of flight capability and included measures of wing span, area, aspect ratio and the second moment of wing area.
3. Three distinctly different distribution patterns of mature female caddisflies were identified. The abundance of three species varied over larger scales only (multiple channel units). Six species that oviposit preferentially in riffles had higher female abundances at riffles than pools, but for only one did abundances also vary over larger scales. There was no association between these different patterns and measures of wing morphology, after removing metrics that were correlated and that differed systematically between taxonomic families. However, we could not reject the hypothesis that some aspect of flight behaviour may have contributed to observed patterns.
4. The diverse but distinct distributions of mature female caddisflies we observed along short channel lengths are novel and suggest that species differ in their propensity for movement along streams, which could have consequences for local densities of eggs and juveniles in the aquatic environment. The degree to which population sizes are coupled across the terrestrial-to-aquatic transition is rarely investigated in aquatic insects and may provide fresh insight into sources of spatial variation within populations. Similarly, a more nuanced approach to research on the flight of aquatic insects, including age- and sex-specific phenomena, may provide greater insight into the diverse ecological functions and consequences of movement.

     

Do pools impede drift dispersal by stream insects?

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The relation between dispersal and survival of Lobesia botrana larvae and their density in vine inflorescences

Dispersal and survival of Lobesia botrana Den. & Schiff. (Lepidoptera: Tortricidae) larvae in a simulated first generation and the relationship with their density on vine inflorescences were studied under field conditions. Artificial infestations with neonate larvae were conducted at densities of 5, 10, 15, 20 and 25 individuals per vine inflorescence. Larvae had a considerable dispersal capacity on the vine espalier and were able to reach several inflorescences around those artificially infested. Dispersal downwards (63.1%) was significantly more frequent than upwards (36.9%), probably because larvae move down the vine plant using silk threads. However, the fact that there was upwards dispersal provides evidence that larvae are capable of active locomotion upwards on the vine plant structure. Mean distances covered by larvae ranged between 10 and 30 cm, with a maximum as far as 45 cm. The longest displacements were not associated with the downward dispersal. The maximum distance covered by larvae was positively correlated with larval density. Mean distance and larval density were not correlated, but mean distances covered at high larval densities were significantly higher than at low densities. At higher larval densities, the proportion of larvae which established in the artificially infested inflorescences decreased whereas the proportion of dispersing individuals increased. However, as a result of the balance between establishment and dispersal, larval survival did not differ significantly among larval densities (26–44%). The results obtained suggest that larval dispersal must be taken into account when preimaginal stages are sampled to determine whether damage thresholds are reached in an integrated pest management program.     

Predation on mayfly nymph, Baetis rhodani, by native and introduced Gammarus: direct effects and the facilitation of predation by salmonids

1. In a series of laboratory experiments, we assessed the predatory nature of the native Irish amphipod, Gammarus duebeni celticus , and the introduced G. pulex , towards the mayfly nymph Baetis rhodani . We also investigated alterations in microhabitat use and drift behaviour of B . rhodani in the presence of Gammarus , and indirect predatory interactions with juvenile Atlantic salmon, Salmo salar .
2. In trials with single predators and prey, B. rhodani survival was significantly lower when Gammarus were free to interact with nymphs as than when Gammarus were isolated from them. The invader G. pulex reduced the survival of B. rhodani more rapidly than did the native G. d. celticus . Both Gammarus spp. were active predators.
3. In `patch' experiments, B. rhodani survival was significantly lower both when G. pulex and G. d. celticus were present, although the effect of the two Gammarus species did not differ. Again, active predation of nymphs by Gammarus was observed. Significantly more nymphs occurred on the top and sides of a tile, and per capita drifts were significantly higher, when Gammarus were present. Baetis rhodani per capita drift was also significantly higher in the presence of the introduced G. pulex than with the native G. d. celticus .
4.  Gammarus facilitated predation by salmon parr of B. rhodani by significantly increasing fish–nymph encounters on exposed gravel and in the drift. There were no differential effects of the two Gammarus spp. on fish – B . rhodani encounters or consumption.
5. We conclude that Gammarus as a predator can have lethal, nonlethal, direct and indirect effects in freshwaters. We stress the need for recognition of this predatory role when assigning Gammarus spp. to a `Functional Feeding Group'.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

Life histories, microdistribution and drift of two mayfly (Ephemeroptera) species in the Pembina River, Alberta, Canada

Ephemerella inermis Eaton is a univoltine species that emerges in July. Eggs hatch in August and larvae grow rapidly in autumn and spring, but not during winter. Baetis tricaudatus Dodds produces three cohorts per year. Emergence periods occur in early June, July and early September. Only one cohort overwinters as larvae.
Microdistribution of both species is controlled largely by the discharge pattern of the river. During 1978, a year of frequent floods, animals were equally distributed among slow and fast water portions of the river. Nocturnal drift densities of both species were much greater in slow than in fast water areas. During 1979, a year of relatively stable flow, benthic larval densities were significantly greater in fast water than in slow water.
Early instar E. inermis larvae are most abundant in nearshore areas, but move to deeper water in late autumn. Baetis tricaudatus larvae are seldom numerous in near-shore areas.
Densities of both species were positively associated with concentrations of detritus and of other invertebrates, but only when relationships were considered in areas of similar current velocity independently of season.     

Recruitment, kin and the spatial genetic structure of a caddisfly Plectrocnemia conspersa in a southern English stream

1. Our objectives were to examine small‐scale patterns in oviposition and genetic relatedness in a population of the stream‐dwelling caddis Plectrocnemia conspersa and, in particular, to look for any evidence of the spatial proximity of close kin and, therefore, ‘patchy recruitment’. 2. In order to examine the distribution of related larvae at the beginning of the aquatic phase, we searched the stream for egg masses. Spatially and temporally structured samples of larvae were then collected from the stream over four sample dates within one generation. The genetic relatedness of these field‐collected larvae, estimated using six polymorphic microsatellite loci, was subsequently compared with that of larvae reared from individual egg masses in the laboratory. 3. Egg masses were laid in clusters or ‘hot spots’ along the stream. The mean relatedness coefficient within reared egg masses ranged from 0.327 to 0.525, and differed significantly from that estimated for the population as a whole (0.070), indicating that the markers were sufficiently powerful to identify groups of siblings. Each egg mass was likely to be the progeny of one father, although the possibility of a small contribution from a second male could not be excluded for a few masses. 4. Mean relatedness within the spatially structured groups of larvae in the stream, even those in close proximity to each other, did not differ from the background population level estimate, suggesting that siblings disperse away from each other very quickly and that kin structure does not persist over time. 5. Changes in spatial genetic structure late in the larval life indicated that neighbouring larvae were less closely related than the population overall, possibly suggesting some direct or indirect means of avoiding kin when approaching the onset of pupation. 6. Our counts of egg masses suggested that survival through the egg stage and early larval life was apparently very high (>50%) for a non‐social insect, and may be a consequence of the colonial net that is briefly occupied by first instar larvae. 7. The number of egg masses laid in the stream (approximately 400 in the sample year), the lack of a spatial genetic structure showing evidence of families even early in larval life, and the high survival of eggs and early instar larvae, all refute the ‘patchy recruitment hypothesis’ for this species.     

Drift and settlement of stream insects in a complex hydraulic environment

1. The hydraulic and geomorphic characteristics of stream patches are often associated with distinctive assemblages or densities of stream invertebrates, and it is routinely presumed that these patterns reflect primarily species‐specific habitat requirements. An alternative hypothesis is that such patterns may be influenced by constraints on movement, such as the results of departure and settlement processes. We describe a manipulative experiment that examined how the hydraulic environments created by topographic bedforms influenced the drift behaviour and potential settlement sites for two species of mayfly (Baetis rhodani and Ecdyonurus torrentis). These species are common in the drift and often co‐occur in streams, but differ in their small‐scale distribution patterns, body shape and movement behaviour. 2. Flume experiments were carried out to determine how the hydraulic environments conditioned by a step bedform influence the behaviour of mayflies in the drift (swimming, posturing, tumbling), and the consequences of those behaviours (drift distance and time), compared to drift over a plane bed. The ramped step in the flume mimicked step bedforms that are common in coarse‐grained, high‐gradient streams. In contrast to the plane bed, a zone of recirculating flow was created downstream of the step, above which flow was faster and more turbulent. Uniform flows are used in most flume studies of drift; our approach is novel in recreating a complex hydraulic environment characteristic of stream channels. 3. Both species had some behavioural control over drift, and drift distances and times were shorter for live larvae than for dead larvae over the plane bed. The step had no impact on drift time or distance for live Baetis, but dead larvae were trapped in the flow separation eddy and drift time increased accordingly. Some Ecdyonurus also became trapped in the eddy, but live larvae drifted farther than dead larvae, and farther over the step than the plane bed. 4. Whilst in the drift, larvae altered their behaviour according to the ambient hydraulic environment, but in a species‐specific manner. Over the plane bed, Baetis had occasional swimming bursts, but primarily postured (maintained a stable body orientation), whereas Ecdyonurus spent roughly equal time posturing and swimming. In the more turbulent flows generated by the step, Baetis spent proportionately more time swimming, whereas Ecdyonurus spent more time posturing and often tumbling as body orientation became unstable. 5. In a high‐gradient stream, Baetis was more abundant close to steps than in plane bed patches with less complex flow, whereas the opposite pattern held for Ecdyonurus. Thus, the small‐scale distribution patterns of these species within streams correspond to their drift behaviours and ability to access various hydraulic patch types in our flume. These results are consistent with the hypothesis that constraints on movement and settlement may be important driver of distribution patterns within streams.     

Spatial patterns at host and forest stand scale and population regulation of the pine processionary moth Thaumetopoea pityocampa

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