Biotic and abiotic factors affecting survival of early life history intervals of a stream-dwelling sunfish

Synopsis Biotic interactions, such as competition and predation, and abiotic events, such as floods, both have important consequences for organisms inhabiting warmwater streams. Empirical studies can lead to better understanding of how these factors interact to affect distinct life history intervals of stream fishes. Survival of eggs, embryos and larvae in nests of longear sunfishLepomis megalotis was monitored in a section of Jordan Creek for four years. During two years characterized by low, relatively stable flow, nest failures could be attributed to biotic interactions. During two years with more variable flow, most brood losses occurred during floods. Flooding led to nest desertion and loss of offspring regardless of nest location. The number of nests constructed tended to be lower during years with more variable flow. The proportion of nests with successful broods did not differ between colonial and solitary nests. Variation in flow regime modifies the importance of biotic effects on survival of early life history intervals of longear sunfish.     

Intraspecific variation in group size in the blackbuck antelope: the roles of habitat structure and forage at different spatial scales

The main ecological factors that are hypothesized to explain the striking variation in the size of social groups among large herbivores are habitat structure, predation, and forage abundance and distribution; however, their relative roles in wild populations are not well understood. I combined analyses of ecological correlates of spatial variation in group size with analyses of individual behaviour in groups of different sizes to investigate factors maintaining variation in group size in an Indian antelope, the blackbuck Antilope cervicapra. I measured group size, habitat structure, forage, and the occurrence of predators in ten blackbuck populations, and, at a smaller spatial scale, within an intensively studied population. To examine the processes by which these ecological factors influence group size, I used behavioural observations and an experiment to estimate the shape of the relationship between group size and potential costs and benefits to individuals. Group size varied extensively both among and within populations. Analyses of spatial variation in group size suggested that both forage and habitat structure influence group size: large-scale, among-population variation in group size was primarily related to habitat structure, while small-scale, within-population variation was most closely related to forage abundance. Analyses of individual behaviour suggested that larger groups incur greater travel costs while foraging. However, individuals in larger groups appeared to experience greater benefits, namely the earlier detection of a “predator”, a reduction in vigilance, and an increase in the time spent feeding. Overall, these findings suggest that individuals in groups experience a trade-off between predation-related benefits and costs arising from feeding competition. Habitat structure and forage likely influence the nature of this trade-off; thus, variation in these ecological factors may maintain variation in group size. The role of predation pressure and other factors in explaining the remaining variation needs further exploration. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Reedbed structure and habitat preference of reed passerines during the post-breeding period

Post-breeding habitats of nine passerine species were studied in the permanently flooded reed beds of ponds near the village Pusté úlany (SW Slovakia) in 2002. Structural features of reed beds and the abundance of all birds studied were sampled at two sites and eight study plots. The reed bed structure and abundance of four passerines differed considerably between the individual study plots. The variations in vegetation structure were also caused by winter reed burning in the SE part of study area. Stems in burnt reed beds were smaller and thinner than unburnt. Abundance of the reed warbler and to a lesser extent sedge warbler was higher in tall reed beds, while the bearded tit preferred thick reed stems. Position of reed beds along the shore — open water gradient was an important factor in terms of abundance at the study plots in the case of the reed bunting. This species showed a strong preference for the reed bed edge with open water. Horizontal distribution of other species seems to be random, however, the birds with the exception of Savi’s warbler preferred the upper stratum of reed bed in vertical distribution. Our field data corroborate with those found in experimental studies concerning with ecomorphological characteristics of birds. Some inconsistencies, however, could also be caused by other factors (e.g. foraging preferences). Comparison of breeding and post-breeding habitat requirements of the studied bird species confirmed a more or less random distribution of the majority of species in the post-breeding period. Some variation was found in the cases of only the reed warbler and bearded tit.     

Population niche structure

Summary The response of twenty maternal families of the annual Abutilon theophrasti to two resource gradients, nutrient and light, was investigated. The structure of the population niche for both biomass and reproductive output was found to be quite different on the two gradients. On the light gradient there was a great diversity of responses among the families while on the nutrient gradient the families responded in a similar manner. On both gradients the plants showed a significant genotype/environment interaction. Three strategies for the production of seed variation have been proposed-all offspring are adapted to the same restricted environment, each offspring of an individual is adapted to a particular environment somewhat different thant that of its siblings, and all the offspring are able to grow in a wide range of environments. We found evidence for all three of these strategies amongst the families. The range of responses seen amongst families (of the same species) in this study was as broad as that found in previous studies among species of the old field annual community to which Abutilon theophrasti belongs. This has significant implications to the nature of competitive interactions and to the evolution of differential resource use in plant populations.     

Seagrass beds and intertidal invertebrates: an experimental test of the role of habitat structure

The majority of field experiments have been carried out on relatively small spatial and short temporal scales, but some of the most interesting ecological processes operate at much larger scales. However, large-scale experiments appropriate to the landscape, often have to be carried out with minimal plot replication and hence reduced statistical power. Here, we report the results of such a large scale, un-replicated field experiment on the Mondego estuary, Portugal, which nevertheless provides compelling evidence of the importance of habitat structure for invertebrate community composition and dynamics. In this estuary, seagrass beds have suffered a dramatic decline over the last 20 years, associated with changes in invertebrate assemblages. In addition, the most abundant species in the system, Hydrobia ulvae, displays distinctly different population structures in those sites. The aim of the field experiment was to test the hypothesis that these differences are related to enhanced survival of snails due to protection from avian or fish predators so that they can grow to larger body sizes in the more complex habitat provided by seagrass. We tested this hypothesis through a large-scale experiment using artificial seagrass beds over a 12-month period. Adult snail densities were higher in the artificial bed plots compared to controls. However, these differences emerged only slowly, related to snail growth rate. This suggests that protection from epibenthic predators can have a significant effect on population structure and hence biomass and productivity of key species in this system. However, the invertebrate assemblage in artificial seagrass plots and the natural seagrass bed, remained statistically separate by the end of the experiment. Handling editor: K. Martens     

Response of harpacticoid copepods to habitat structure at a deep-sea site

Given the quiescent physical environment and the low rates of bioturbation in the deep sea, biologically produced structures, such as animals tubes, may play an important role in creating habitat heterogeneity. In San Diego Trough, the cirratulid polychaete Tharyx luticastellus builds and inhabits a robust mud concretion (a mudball). Mudballs are abundant and persist after the worm has died or abandoned them. Our analysis of twelve 6.6-cm-diameter cores from 32° 51.02 N, 117° 46.97 W, taken with the submersible SEA CLIFF, indicate that one of 29 species of harpacticoids common in San Diego Trough is associated with Tharyx mudballs. This species (of the genus Heteropsyllus) responded only to mudballs containing worms, suggesting that any effects of the inanimate structure were not the cause of the association. We could detect no difference in bacterial numbers in sediments between control and occupied-mudball samples. This result suggests that the response does not result from gross differences in food levels near a mudball, but the possibility that there are differences in some preferred food cannot be discounted. An univestigated possibility is that Heteropsyllus responds to occupied mudballs because the worm's presence deters some (presumably small) predator.     

Quantifying microhabitat availability: stratified random versus constrained focal-fish methods

During a study of microhabitat use by gilt darters (Percina evides), we compared two methods for quantifying microhabitat availability in a southern Appalachian stream (USA). The first method used stratified random sampling throughout the site and the second involved taking constrained random measurements within a 2-m radius of the focal fish. Darters were generally over-represented in microhabitats with higher average velocities, greater amounts of erosional substrata, and lower amounts of depositional and large substrata. The two methods generally yielded similar patterns of microhabitat use. Nonetheless, of the seven microhabitat categories in which differential microhabitat use occurred in summer, four were present in both data sets, but three differed between methods. We observed no differences between methods for autumn data. Finally, the standard deviations of the summer-stratified random data set were significantly greater (sign test, P < 0.05) than those of the constrained data set. Our results suggest that either method for quantifying microhabitat availability can be used to quantify the general habitat use patterns of this species, but constrained analyses yielded a more restricted view of the total habitat available. Nonetheless, if the fishes range over a site, clearly stratified habitat availability analysis is preferred. Handling editor: J. A. Cambray     

Trophic interactions and habitat segregation between competing Daphnia species

Summary Two commonly coexisting species of Daphnia segregate by habitat in many stratified lakes. Daphnia pulicaria is mostly found in the hypolimnion whereas D. galeata mendotae undergoes diel vertical migration between the hypolimnion and the epilimnion. I examined how habitat segregation between these two potentially competing species might be affected by trophic interactions with their resources and predators by performing a field experiment in deep enclosures in which I manipulated fish predation, nutrient levels, and the density of epilimnetic Daphnia. The results of the experiment indicate that habitat use by D. pulicaria can be jointly regulated by competition for food from epilimnetic Daphnia and predation by fishes. Patterns of habitat segregation between the two Daphnia species were determined by predation by fish but not by nutrient levels: The removal of epilimnetic fish predators resulted in higher zooplankton and lower epilimnetic phytoplankton densities and allowed D. pulicaria to expand its habitat distribution into the epilimnion. In contrast, increased resource productivity resulted in higher densities of both Daphnia species but did not affect phytoplankton levels or habitat use by Daphnia. The two species exhibit a trade-off in their ability to exploit resources and their susceptibility to predation by fish. D. g. mendotae (the less susceptible species) may thus restrict D. pulicaria (the better resource exploiter) from the epilimnion when fish are common due to lower minimum resource requirements than those needed by D. pulicaria to offset the higher mortality rate imposed by selective epilimnetic fish predators. D. g. mendotae does not appear to have this effect in the absence of fish.     

The effects of spatial habitat structure on population variability of freshwater snails

Experiments were conducted to examine the effects of a habitat's spatial structure on population variability in two species of freshwater snails (Physa acuta and Austropeplea ollula). To alter the spatial structure of the habitat, vinyl chloride plates were hung in experimental tanks, providing three types of spatial structure: Complex structure, Simple structure and Control (no structure). In Experiment 1, the average number of individuals in a tank did not differ among the three types of structure 2 months after the introduction of the snails, but the variability of the number of individuals in the Complex structure tanks was lowest, whereas the variability in the Control tanks was highest. In Experiment 2, in addition to the spatial structure of the habitat, three types of species interaction were designed as experimental treatments: only P. acuta was introduced into the tanks (P. acuta tanks), only A. ollula was introduced into the tanks (A. ollulatanks) and both P. acuta and A. ollula were introduced into the tanks (two-species tanks). For the P. acuta tanks, the variability of the number of P. acuta individuals in the Complex structure tanks was lowest, and the variability in the Control tanks was highest when the effect of the number of individuals in a tank was subtracted. For the A. ollula tanks and the two-species tanks, there were no significant differences in the variability of the population size among the different treatments of spatial structure. The spatial distribution of P. acuta was more uniform than the distribution of A. ollula on the plates of complex structure. Our results indicate that the spatial structure of the habitat influences the variability of population size (the variance of the number of individuals in different populations during the earlier period after the introduction of the snails), but the effects depend on the spatial behavior of individuals and the interaction with other species.     

Posture, speed, and habitat structure: three-dimensional hindlimb kinematics of two species of padless geckos

Differences in habitat use are often correlated with differences in morphology and behavior, while animals in similar habitats often exhibit similarities in form and function. However, this has not been tested extensively among lizards, especially geckos. Most studies of gecko locomotion have focused on the ability to adhere to surfaces. However, there are several species of geckos that have either secondarily lost adhesive capabilities or simply lack the capability. We quantified the three-dimensional locomotor kinematics for two desert-dwelling padless geckos, Teratoscincus scincus and Eublepharis macularius, on a level trackway over a range of speeds. Our results indicate that T. scincus landed with a high relative hip height of 48.7 ± 2.4% of total limb length at footfall, while E. macularius exhibited hip heights averaging only 36.0 ± 1.8% of total limb length for footfall. The three-dimensional knee angle of T. scincus averaged 120.6 ± 3.9° at footfall, while E. macularius averaged only 101.6 ± 1.8° at footfall. In addition, the femur of E. macularius was elevated to a much greater extent (i.e., was closer to being perpendicular to the long axis of the body) than that of T. scincus and every other lizard that has been studied, suggesting they move with a “hyper-sprawled” posture. Both of these gecko species live in deserts, but T. scincus is psammophilic while E. macularius inhabits a rocky, more densely vegetated environment. Benefits of the more upright posture of T. scincus on open sandy habitat may include a greater field of view and more efficient locomotion. The more sprawled posture of E. macularius may lower its center of gravity and aid in balance while climbing on rocks or shrubs.     

Reproductive ecology of red-chinned lizards (Sceloporus undulatus erythrocheilus) in Southcentral Colorado: comparisons with other populations of a wide-ranging species

Summary We studied the reproductive ecology of a population of Sceloporus undulatus erythrocheilus near Walsenburg, CO during the summers of 1987–1990. Reproductive activity commences soon after emergence in early May and continues until the middle of July. Females mature in their second year following hatching at an age of 20–21 months and a size of 60–63 mm snout-vent length (SVL). Two clutches, averaging10.9 eggs, are produced per year. Larger females produce larger clutches with an increase of one egg per 3 mm SVL. Average SVL of females was 71 mm. Eggs are reasonably large (0.32 g) and relative clutch mass was the largest reported for the species (34% of body mass). A principal component analysis was used to examine relationships among reproductive characters in 12 populations of Sceloporus undulatus. The analysis suggested that patterns of reproduction in this wide-ranging species result from several factors reflecting both adaptive and phylogenetic sources of variation.     

Loud calls ofGalago crassicaudatus andG. garnettii and their relation to habitat structure

Long distance vocalizations have been shown to be good indicators of genetic species in primates. Here the loud calls of two recently identified greater galago taxa —Galago crassicaudatus andG. garnettii — are compared and analyzed statistically. Observed differences in call structures are investigated further as potential indicators of differences in the structures of habitats frequented by the two species. Although the calls share a repetitive structure, and show similar dominant frequency bands (1,000 – 1,500 Hz), they differ significantly in the number of units per call, unit duration, inter-unit interval, highest frequency, lowest frequency, dominant frequency band, first harmonic, and call duration. The duration of theG. crassicaudatus call is more than twice that ofG. garnettii. Strong intraspecific consistency is seen in the most energetic frequency bands (dominant frequency band and first harmonic), and durations of the individual units and inter-unit intervals. Information important to species recognition is thus most likely to be contained in these features. Individual recognition may be encoded in the relative emphasis of higher level harmonics. The frequency structures of the calls will reflect requirements for acoustical transmission in a forest environment, as well as structural constraints imposed by body size. Higher frequencies detected in theG. garnettii call (up to 8,500 Hz) may have a functional significance related to distance estimation, or may simply be a reflection of smaller body size. The greater modulation of theG. garnettii call suggests that its habitat constitutes a denser or more turbulent medium for sound transmission than does the habitat ofG. crassicaudatus.     

Large- and small-scale effects of habitat structure on rates of predation: how percent coverage of seagrass affects rates of predation and siphon nipping on an infaunal bivalve

Landscape ecology, predominantly a terrestrial discipline, considers the effect of large-scale (tens of meters to kilometers) spatial patterns of habitats on ecological processes such as competition, predation, and flow of energy. In this study, a landscape-ecology approach was applied to a marine soft-sediment environment to examine rates of predation and transfer of secondary production in and around vegetated habitats. Seagrass beds naturally occur in a variety of spatial configurations from patches 1–10s of meters across with interspersed unvegetated sediments (i.e., patchy coverage) to more continuous coverage with little or no bare sediment. I designed experiments to address how percent coverage of seagrass in a 100-m² area of seafloor, and the spatial arrangement (degree of patchiness or fragmentation) of an equal area (100 m²) of vegetation affected predation (lethal) and siphon nipping (sublethal) intensity on an infaunal bivalve, Mercenaria mercenaria (hard clam). Measures of seagrass density and biomass with different percent coverage of seagrass were also made. When clams were placed in both the vegetated and unvegetated portions of the seafloor nearly twice as many clams were recovered live with 99% seagrass cover than with 23% seagrass cover, while survivorship was intermediate with 70% cover. Cropping of clam siphons from both the vegetated and unvegetated sediments was also affected by the amount of seagrass cover in a 100-m² area of seafloor: mean adjusted siphon weights were approximately 76% heavier from the 99% seagrass cover treatment than from the 70% or 23% cover treatments. Survivorship of clams placed within an equal area of seagrass in very patchy, patchy, and continuous spatial configurations was 40% higher in the continuous seagrass treatment than in either of the two patchy treatments. This study demonstrates that transfer of secondary production in the form of predation and cropping on an infaunal organism is altered as the percent cover of seagrass changes. While large-scale changes in the amount and spatial patterning of vegetation may affect habitat utilization patterns and foraging HGLoopbehavior, increased seagrass density and biomass with increased percent coverage of seagrass limit any conclusions concerning predator foraging behavior and feeding success in response to patch shapes and sizes. Instead, local changes in seagrass characteristics provide the most compelling explanation for the observed results.     

Isozyme variation and correspondence with unusual infrared reflectance patterns inPhragmites australis (Poaceae)

Isozyme variation was investigated in populations ofPhragmites australis (Poaceae) which have recently invaded and taken over marsh habitat of the Mississippi River delta. Infrared aerial photographs in the Garden Island Bay subdelta of the extensiveP. australis populations reveal distinct, clone-like circular patches within a predominant background. Preliminary evidence indicates that the infrared color differences represent distinct morphological types. However, there are no obvious environmental factors that could account for the peculiar patterns.P. australis collections were taken from five separate and distinct patches and adjacent background. Only two electrophoretic phenotypes were found: one from patches and one from the background. In comparing the two, 20% of the 40 loci scored are fixed for alternate alleles. These results indicate a clear correspondence of infrared reflectance with electrophoretic phenotype. In addition, the genetic uniformity as evidenced by the discovery of only two electrophoretic phenotypes supports the contention that the recent spread ofP. australis throughout the Mississippi River delta has been primarily, if not exclusively, a result of vegetative propagation.     

Population structure and mycelial phenotypic variability of the ectomycorrhizal basidiomycete Laccaria bicolor (Maire) Orton

Mating type allele distribution and phenotypic variability were investigated in field populations of Laccaria bicolor. Sporophores associated with Norway spruce (Picea abies), colonized by natural sources of inoculum and growing in a seed orchard, were sampled to obtain dikaryotic strains and assay their phenotypic variability for traits important to the symbiosis. Basid-iospores were also collected for mating type analysis of different mycelia. Four sporophore mating types were identified containing seven A and five B factors. Out-breeding efficiency was estimated at 73.8% and the population was slightly inbred. Crosses with previously characterized L. bicolor strains from two nearby populations identified in total six sporophore mating types and ten A and nine B factors, for an estimated outbreeding efficiency (85.7%) similar to previous studies of more spatially disparate Laccaria spp. populations. Dikaryotic strains were tested for mycelial growth rate, as a measure of their competitive ability, on agar media containing a soluble (NaH₂PO₄), or an insoluble (CaHPO₄) phosphate source. Their ability to solubilize the latter was also tested to assess their relative capacity to access insoluble, inorganic phosphate. In most cases, significant variation was detected among strains from the same site for all variables. On three sites (VC4, VC5 and VC7), each determined previously to possess a uniform mycelial genotype, phenotypic variability was likely due to epigenetic variation among different strains of the same genotype. Possible evidence for dikaryon-monokaryon crosses was observed in vivo on one sample site (VC2) where adjacent mycelia shared two mating factors. The phenotypic variability of different mycelial genotypes reflected their genetic variability observed as mating type allele diversity and underlined the importance of basidiospore dispersal in introducing new genotypes into the population. The reproductive strategies of L. bicolor are discussed and compared to those of other basidiomycete species.     

Effect of habitat structure on positional behavior and support use in three species of lemur

This study compares locomotor and postural behavior and substrate use of three species of lemur, the diademed sifaka (Propithecus diadema edwardsi), the brown lemur (Eulemur fulvus rufus), and the red-bellied lemur (Eulemur rubriventer) at two different localities within Ranomafana National Park. The object of the study is to see if there are quantitative differences in the behavior of the lemurs or their choice of substrates in forests that have different structural attributes. Analysis of the physical characteristics of the habitat demonstrates that compared to the Talatakely area, the forest at Vatoharanana has a higher proportion of larger, taller trees. The behavior of the lemurs also differs in the two areas: all species leap less and climb and move quadrupedally more at Vatoharanana. All species use small size supports less frequently at Vatoharanana, choosing insted medium size supports (all three species) or tiny supports (Propithecus andE. fulvus) found in tree crowns and terminal branches. The lemurs prefer (i.e. use more often than would be expected based on abundance) large trees at both sites. At Vatoharanana however, they are more frequently observed higher in trees and in taller trees with greater trunk breadth. The differences in locomotor behavior are in part due to the fact that at Vatoharanana, more bouts are collected during feeding and foraging than during travel. The relationship between this difference in activity pattern and the structural differerences in the two forests, however, is not clear. This study points out the need for longer term field studies of positional behavior and substrate use that incorporate the variety of forest types the subject species inhabit.     

Population structure ofSclerotium rolfsii in peanut fields

Sclerotium rolfsii isolates from peanut fields in Ibaraki were classified into mycelial compatibility groups (MCGs) based on the barrage zone formation. A total of 132 isolates collected from four fields within a 120 m radius in 1994 comprised four MCGs; MCG A (71 isolates), B (34 isolates), C (26 isolates) and D (one isolate). Fields 1 and 2 were occupied exclusively by MCG A. MCG A also predominated in field 3. In field 4, MCGs A, B and C were dominant. Population structure in 3 additional fields was determined in 1997. All 11 isolates from Field 5, which was 400 m distant from field 1, belonged to MCG C. A total of 42 isolates from fields 6 and 7, 2.5 km distant from other fields and 100 m distant from each other, were all MCG A. These results suggested that the population structure ofS. rolfsii was simple. RAPD fingerprintings showed that most isolates of the same MCG were clonal.     

Population structure and responses to disturbance of the basidiomycete Resinicium bicolor

Summary Resinicium bicolor (Alb. & Schw. ex Fr.) Parm. [=Odontia bicolor (Alb. & Schw. ex Fr.) Bres.] is an outcrossing resupinate basidiomycete associated with root and butt rots of trees, but is itself only very weakly pathogenic. The distribution of genets among every spruce stump in a 70-year-old 1250 m² spruce stand was analysed using somatic incompatibility testing. R. bicolor was present on 40% of 8-to 10-year-old stumps. Nineteen genets were found occupying 32 stumps; yielding probabilities of colonisation following establishment by basidiospores of 0.20–0.24 and by mycelial extension or dispersal of 0.16–0.20. The probability of colonisation decreased with increasing distance from a point of establishment. R. bicolor responded to both enrichment and destructive disturbances by the formation of an extensive cord system which enabled it to colonise discontinuously distributed resources and to overgrow fungi adjacent to it in a single resource unit, including Heterobasidion annosum.     

Individual food niche specialization in stream-dwelling charr

Food niche variation within a population of white-spotted charr, Salvelinus leucomaenis, was investigated by means of mark and recapture to examine the simplification that conspecific individuals are ecologically equivalent. Food diversity measured on the basis of living form of dietary organisms demonstrated variability in food niche in incidence, degree, and stability. As inferred from body size distribution and fish movement, niche variation was inexplicable by social rank hierarchy or foraging habitat rotation, and occasional niches were deemed to be realized individually. Higher annual growth rate shown by those with a larger niche width indicated the adaptive significance in niche variation. Assuming trade-off between feeding efficiency and predation risk, phenotypic plasticity was suggested to determine the food niche specialization of individuals to maximize lifetime fitness.