Predation hazard and seed removal by small mammals: microhabitat versus patch scale effects

Predator avoidance may involve response strategies of prey species that are time and space specific. Many studies have shown that foraging individuals avoid predators by altering microhabitat usage; alternatively, sites may be selected according to larger-scale features of the habitat mosaic. We measured seed removal by two small mammal species (Peromyscus leucopus, and Microtus pennsylvanicus) at 474 stations over an experimentally created landscape of 12 patches, and under conditions of relatively high (full moon) and low (new moon) predatory hazard. Our objective was to determine whether predator avoidance involved the selection of small-, medium-, or large-scale features of the landscape (i.e., at the scale of microhabitats, habitats, or habitat patches). We found rates of seed removal to vary more with features of whole patches than according to variation in structural microhabitats within patches. Specific responses included: under-utilization of patch edge habitats during full moon periods, and microhabitat effects that were only significant when considered in conjunction with larger-scale features of the landscape. Individuals residing on large patches altered use of microhabitats/habitats to a greater extent than those on smaller patches. Studies just focusing on patterns of microhabitat use will miss responses at the larger scales, and may underestimate the importance of predation to animal foraging behavior.     

Behavior of fish predators and their prey: habitat choice between open water and dense vegetation

Synopsis Behavior of largemouth bass, Micropterus salmoides, and northern pike, Esox lucius, foraging on fathead minnows, Pimephales promelas, or bluegills, Lepomis macrochirus, was quantified in pools with 50% cover (half the pool had artificial stems at a density of 1000 stems m⁻²). Both predators spent most of their time in the vegetation. Largemouth bass searched for bluegills and ambushed minnows, whereas the relatively immobile northern pike ambushed all prey. Minnows were closer to predators and were captured more frequently than bluegills. Even when minnows dispersed, they moved continually and eventually wandered within striking distance of a predator. Bluegills dispersed in the cover with predators. Bass captured the few bluegills that strayed into the open and pike captured those that approached too closely in the cover. The ability of predators to capture prey while residing in habitats containing patches of dense cover may explain their residence in areas often considered to be poor ones for foraging. The unit is sponsored jointly by the United States Fish and Wildlife Service, Ohio Department of NaturalResources, The Ohio State University, and the Wildlife Management Institute     

Interactions among coexisting larval Odonata: an in situ experiment using small enclosures

Field experiments using small replicated enclosures focused on interactions between larval populations of Epitheca cynosura and Ladona deplanata (Odonata: Anisoptera) — two species that emerge in early spring. The presence of Epitheca reduced the total biomass of Ladona, but Ladona had no significant effect on Epitheca. These early-emerging species reduced the biomass of small instars of late-emerging Anisoptera which colonized enclosures during the experiments; and the late-emerging Anisoptera seem to have inhibited colonization by Zygoptera larvae. Results are consistent with the importance of predatory (cannibalism or mutual predation) interactions in this community.     

Microhabitat selection by larval Chironomus tentans (Diptera: Chironomidae): effects of predators, food, cover and light

1. As part of a study designed to estimate the developmental costs of antipredator behaviour of larval chironomids, we used laboratory experiments to study effects of food and factors that could influence predation risk [presence of fish, cover from fish (simulated debris) and light level], on microhabitat selection by Chironomus tentans larvae in the third and fourth instar. 2. Larvae were more likely to build tubes where there was more food although their ability to move far to find food appeared limited. 3. Larvae did not avoid areas with fish and the presence of fish did not alter larval response to food. 4. Larvae avoided areas of cover (simulated debris) but cover did not alter larval response to food. 5. When provided with a choice between light and dark areas, larvae initially without tubes were found more often in the dark areas. Light level had no effect on location of larvae that had begun the experiment with tubes. 6. Results suggest the tubicolous life-style of larval Chironomus tentans limits their ability to select microhabitats that could alter their risk of predation.     

Behavioural Response of Juvenile Bluegill Sunfish to Variation in Predation Risk and Food Level

The behavioural response of juvenile bluegill sunfish (Lepomis macrochirus) to predation risk when selecting between patches of artificial vegetation differing in food and stem density was investigated. Bluegill foraging activity was significantly affected by all three factors. Regardless of patch stem density or risk of predation bluegills preferred patches with the highest prey number. During each trial bluegill foraging activity was clearly divided into a between- and within-patch component. In the presence of a predator bluegills reduced their between-patch foraging activity by an equivalent amount regardless of patch stem density or food level, apparently showing a risk-adjusting behavioural response to predation risk. Within patches, however, foraging activity was affected by both food level and patch stem density. When foraging in a patch offering a refuge from predation, the presence of a predator had no effect on bluegill foraging activity within this patch. However, if foraging in a patch with only limited refuge potential, bluegill foraging activity was reduced significantly in the presence of a predator. Further, this reduction was significantly greater if the patch contained a low versus a high food level, indicating a risk-balancing response to predation with respect to within-patch foraging activity. Both these responses differ from the risk-avoidance response to predation demonstrated by juvenile bluegills when selecting among habitats. Therefore, our results demonstrate the flexibility of juvenile bluegill foraging behaviour.     

Comparative microhabitat use of cyprinid larvae and juveniles in a lotic floodplain channel

Synopsis The microhabitat of six cyprinids in a braided side-channel of the Upper Rhône River, France, was studied June to September 1985, using point abundance sampling by electrofishing. Correspondence analysis of the samples-by-species data and Jacob&s electivity index (calculated from the availability and exploitation of eight environmental variables) revealed three groups of species: (1) Leuciscus cephalus and Alburnus alburnus, which used relatively similar microhabitat during both larval and 0 + juvenile development; (2) Rutilus rutilus, Leuciscus leuciscus and Chondrostoma nasus, which used different microhabitats as larvae but whose microhabitat overlapped notably during 0 + juvenile development; (3) and Gobio gobio, whose juvenile microhabitat overlapped only marginally with that of other 0 + juveniles (no data were available on larvae). Overlap in microhabitat use by 0 + juveniles increased notably during a period of reduced discharge, when the amount of available vegetal and ligneous structures decreased; while most species exploited the increased area of shallow waters, juvenile A. alburnus appeared to be the only species that exploited the deeper waters off the channel&s steeper banks. Although not quantified, numerous field observations of predation threat, particularly during the period of reduced discharge (which reduced the amount of cover), suggest that predation risk may influence the microhabitat use of young fishes. Compared with similar species in European lakes, the overlap in microhabitat use of these fluvial cyprinids appears to be more intensive.     

Lack of preference for low-predation-risk habitats in larval damselflies explained by costs of intraspecific interactions

Many studies indicate prey organisms select microhabitats with high structural complexity as a way of reducing risk of predation. We used laboratory experiments to show that damselfly larvae, Ischnura verticalis, suffer higher predation rates from pumpkinseed sunfish in low-density vegetation. However, larvae do not preferentially occupy microhabitats with high vegetation density in either the presence or absence of sunfish; when given a choice, the number of larvae per stem of vegetation was equal across all densities of vegetation. That larvae do not congregate in dense vegetation may reflect costs of aggressive interactions. Results from laboratory experiments indicated larval interactions increase conspicuous behaviours (most notably swimming) and consequently increase fish predation. A subsequent experiment indicated that frequency of larval interactions increases with increased vegetation density when number of larvae/stem is constant. Thus, larval microhabitat selection may reflect a trade-off between reduced risk of predation in areas of high vegetation density, caused by reduced fish foraging ability, and increased aggressive larval interactions, due to decreased proximity of larvae. Copyright 2000 The Association for the Study of Animal Behaviour.     

Intraguild predation and interference competition on the endangered dragonfly <Emphasis Type="Italic">Aeshna viridis</Emphasis>

We examined the effects of intraguild predation (IGP) and interference competition on an endangered dragonfly, Aeshna viridis Eversm. (Odonata: Anisoptera). A. viridis is rare in Europe due to the decrease in suitable habitats harboring the macrophyte Stratiotes aloides L. Stratiotes plants are the principal oviposition substrate for A. viridis females and protect the larvae of A. viridis from fish predation. In our study lakes A. viridis larvae are sympatric with larvae of Aeshna grandis and Aeshna juncea. The susceptibility of A. viridis larvae to IGP by similar-sized larvae of A. grandis and A. juncea was tested in a laboratory predation experiment. Microhabitat use of A. viridis and A. grandis was studied in the laboratory to determine the possible effects of interference competition on the spatial distribution of A. viridis larvae. Our results show that at least in laboratory conditions, A. viridis is susceptible to IGP and interference competition. In competition, A. grandis larvae dominated the middle and outer portion of S. aloides rosettes whereas A. viridis stayed in the inner parts. When A. grandis larvae were absent, A. viridis colonized the middle and outer parts of the rosettes. We conclude that asymmetric predation between odonate larvae of equal size can be intense, and that both IGP and interference competition affect A. viridis. Although natural habitat complexity diminishes their impact, these interactions may nevertheless influence the distribution of A. viridis in S. aloides waters and restrict its microhabitat use in S. aloides rosettes.     

The influence of larval case design on vulnerability of Limnephilus frijole (Trichoptera) to predation

• 1 We examined the potential selective pressures exerted by different types of predators on morphological prey defences using the case-building larvae of the limnephilid caddisfly, Limnephilus frijole. The predators were a predaceous minnow, Gila pandora, and an aeschnid dragonfly naiad, Oplonaeschna armata.
• 2 The influence of larval case design on vulnerability to predators was evaluated in a series of laboratory experiments. Larval L. frijole were collected from a stream without fish or dragonflies. In the laboratory, larvae still occupying their field-built cases were exposed to the two predators, with which L. frijole co-occurs in other streams. Following 24 h exposure of caddisfly larvae to a specific predator, the pre-trial case length, case strength, case width, and case composition of victims and survivors were compared.
• 3 Principal components analysis (PCA) indicated substantial independent variation for all four case parameters in the study population. The cases of larvae surviving exposure to either dragonfly naiads or fish were significantly stronger and wider than cases of victims of predation. Cases of survivors were also longer and had a greater mineral fraction than those of victims, but differences were either marginally significant or not significant.
• 4 These results demonstrate that specific aspects of case design affect the vulnerability of caddisfly larvae. In addition, predaceous fish and predaceous invertebrates, using different predation techniques, may exert similar selective pressures on these defenses. Case aspects influencing vulnerability in these experiments are broadly consistent with natural variation in case design associated with predator-rich and predator-free habitats for related caddisfly species.

     

Rodents balancing a variety of risks: invasive fire ants and indirect and direct indicators of predation risk

We used foraging trays to compare how oldfield mice, Peromyscus polionotus, altered foraging in response to the presence of fire ants, Solenopsis invicta, and in the presence of direct (predator urine) and indirect (sheltered or exposed microhabitat, moonlight, and precipitation) indicators of predation risk. Foraging reductions elicited by S. invicta were greater than reductions in response to well-documented indicators of risk (i.e., moonlit nights) and the presence of predator urine. The presence of S. invicta always led to reduced foraging, but the overall impact of S. invicta was dependent upon microhabitat and precipitation. When S. invicta was not present, foraging was greater in sheltered microhabitats compared to exposed microhabitats. S. invicta made sheltered microhabitats equivalent to more risky exposed microhabitats, and this effect was especially pronounced on nights without precipitation. The effect of S. invicta suggests that interactions with S. invicta may entail a potentially heavy cost or that presence of S. invicta may represent a more reliable indicator of imminent competition or predation compared to indirect cues of risk and predator urine. The presence of S. invicta led to reduced foraging under situations when foraging activity would otherwise be greatest (i.e., under vegetative cover), potentially reducing habitat quality for P. polionotus and the distribution of seeds consumed by rodents.     

Habitat-Specific variation in eyespot size of a benthic cladoceran

Summary Pigmented eyespot size of the benthic cladoceran Simocephalus exspinosus was measured in individuals sampled from four freshwater ponds that differed in the extent of visually-oriented predation. In ponds with such predation (from fish, salamander larvae, and dragonfly nymphs), eyespot size was found to be significantly smaller, relative to body size, than in a pond without visually-oriented predation. Reduction in pigmented eyespot size may represent an adaptation to reduce vulnerability to predation.     

Spatial organization of a desert rodent community: food addition and species removal

Summary From 1977 through 1983 we conducted experiments on a desert rodent community where supplemental seeds were added or certain rodent species and ants were removed from 0.25-ha fenced plots in a Chihuahuan Desert site in southeastern Arizona, USA. In this paper we examine the patterns of microhabitat use relative to vegetative cover by 11 rodent species. The results show that: i) removal of the largest seed-eating species, Dipodomys spectabilis, produced the most pervasive and dramatic shifts in microhabitat use by the remaining rodent species; ii) adding seeds or removing ants had little effect on the spatial use of microhabitats by rodents in this community; and iii) non-granivores were just as likely as granivores to shift microhabitat use when other granivores were removed. We believe these results indicate that both food and foraging microsites are limited but the relegation of subdominant species to lesspreferred microhabitats by the large Dipodomys spectabilis is the major factor underlying the spatial organization of this community. Results also demonstrate that strong interactions among species increase the probability that pathways of indirect interactions through intermediary species are important; these complex linkages may include species that overlap little in food preferences.     

Risk of predation and behavioural response in three anuran species: influence of tadpole size and predator type

Many species alter their activity, microhabitat use, morphology and life history in response to predators. Predation risk is related to predator size and palatability of prey among others factors. We analyzed the predation risk of three species of tadpoles that occur in norwestern Patagonia, Argentina: Pleurodema thaul, Pleurodema bufoninum and Rhinella spinulosa. We sampled aquatic insect predators in 18 ponds to determine predator–tadpole assemblage in the study area. In laboratory conditions, we analysed the predation rate imposed by each predator on each tadpole species at different tadpole sizes. Finally, we tested whether tadpoles alter their activity in the presence of chemical and visual cues from predators. Small P. thaul and P. bufoninum tadpoles were the most vulnerable prey species, while small R. spinulosa tadpoles were only consumed by water bugs. Dragonflies and water bugs were the most dangerous tadpole predators. Small P. thaul tadpoles reduced their activity when they were exposed to all predators, while large tadpoles only reduced the activity in the presence of large predators (dragonfly larvae and water bugs). Small P. bufoninum tadpoles reduced the activity when they were exposed to beetle larvae and dragonfly larvae, while large tadpoles only reduced activity when they were exposed to larger predators (water bugs and dragonfly larvae). R. spinulosa tadpoles were the less sensitive to presence of predators, only larger tadpoles responded significantly to dragonfly larvae by reducing their activity. We conclude that behavioural responses of these anuran species were predator-specific and related to the risk imposed by each predator.     

Phenotypic variation and vulnerability to predation in juvenile bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>)

Bluegill sunfish (Lepomis macrochirus) are known to diversify into two forms specialized for foraging on either limnetic or littoral prey. Because juvenile bluegills seek vegetative cover in the presence of largemouth bass (Micropterus salmoides) predators, natural selection should favor the littoral body design at size ranges most vulnerable to predation. Yet within bluegill populations, both limnetic and littoral forms occur where vegetation and predators are present. While adaptive for foraging in different environments, does habitat-linked phenotypic variation also influence predator evasiveness for juvenile bluegills? We evaluate this question by quantifying susceptibility to predation for two groups of morphologically distinct bluegills; a limnetic form characteristic of bluegills inhabiting open water areas (limnetic bluegill) and a littoral form characteristic of bluegills inhabiting dense vegetation (littoral bluegill). In a series of predation trials, we found that bluegill behaviors differed in open water habitat but not in simulated vegetation. In open water habitat, limnetic bluegills formed more dense shoaling aggregations, maintained a larger distance from the predator, and required longer amounts of time to capture than littoral bluegill. When provided with simulated vegetation, largemouth bass spent longer amounts of time pursuing littoral bluegill and captured significantly fewer littoral bluegills than limnetic fish. Hence, morphological and behavioral variation in bluegills was linked to differential susceptibility to predation in open water and vegetated environments. Combined with previous studies, these findings show that morphological and behavioral adaptations enhance both foraging performance and predator evasiveness in different lake habitats.     

The slow—fast life style characteristics in a suite of six species of odonate larvae

• 1 The validity of the slow—fast lifestyle dichotomy proposed by Sih (1987) was tested in a suite of six odonate species from a restricted geographical area. Data on activity and microhabitat use were obtained in a laboratory study. Further necessary information on life history, macrohabitat (ephemeral‐permanent) use and vulnerability to fish predation was provided by a literature survey.
• 2 Activity was estimated as number of moves and distance moved for the six odonate larvae. Aeschna juncea, Lestes sponsa and Sympetrum danae were categorised as high‐active species, whereas Coenagrion hastulatum, Cordulia aenea and Leucorhinia dubia were categorised as low‐active species.
• 3 C. hastulatum and L. sponsa exploited microhabitats close to the water surface, C. aenea and L. dubia close to the bottom, and A. juncea and S. danae were intermediate in their water depth utilisation.
• 4 A principal component analysis of the data from the laboratory experiment and the literature survey supported the slow—fast life style dichotomy since the variables activity, macrohabitat use, life cycle length and sensitivity to fish predators were highly correlated.

     

Plant stem density as a cue in patch choice by foraging juvenile bluegill sunfish

Synopsis Juvenile bluegill sunfish,Lepomis macrochira, are restricted to vegetated habitats by predators. Variation in plant stem density has a significant effect on bluegill foraging success. Given the mosaic nature of this habitat, plant stem density may provide a cue for selecting among patches in which to forage. In this study, juvenile bluegills were offered patches of artificial vegetation differing only in plant stem density as potential foraging sites. Three densities, 100, 250, and 500 stems m^–2 were tested. Fish were presented with a choice between patches (100:250, 250:500, or 100:500). Bluegill foraging rate in, and the number of fish choosing each patch was recorded. Juvenile bluegills showed a preference for those patches which maximized their foraging rate.     

Overdispersion of ant colonies: a test of hypotheses

Summary The nest locations of two ant species in the Colorado Desert are intraspecifically overdispersed. Intraspecific overdispersion has been thought to represent strong intraspecific competition. Here we consider this hypothesis along with three competing hypotheses: microhabitat selection by foundress queens, predation on foundress queens, and predation on established colonies. To test these hypotheses five types of data were collected: (1) the forager population sizes of Veromessor pergandei and Pogonomyrmex californicus, (2) the response of the territory use of V. pergandei to varying levels of food, (3) the encounter rates of conspecifics and other ant species to foundress queens artificially placed near and far from conspecific colonies, (4) predation on colonies as a function of colony spacing, and (5) the relationship between the plant microhabitat at the nest and colony spacing. The results show that established colonies have no apparent selectivity for a particular type of plant microhabitat nor do foundress queens show avoidance or attraction toward conspecifics. V. pergandei workers show only a slight ability to find V. pergandei queens that are artificially placed near their entrances. Certain spiders are the most common ant predators on our study area. Direct observations on spiders indicate that colonies with closer neighbors are not prone to a higher risk of predation. In addition, the estimates of the death rate of workers from a mark-recapture technique indicate that colonies with closer neighbors lose similar numbers of workers as compared to colonies with further neighbors. In favor of the competition hypothesis, the summed size of intraspecific nearest neighbor pairs are larger for colonies that are spaced further apart than those colonies that are spaced closer together. We also develop an index of foraging directionality for the column foraging species V. pergandei. Using this measure, we find that nearest neighbors tend to avoid foraging toward each other. The response of territory use to food levels was tested with experiments involving patches of cracked wheat. These experiments showed that patches away from nearest neighbors were found significantly sooner than patches toward nearest neighbors. The above five sets of data together suggest that resource competition and perhaps queen predation by established colonies account for the intraspecific spatial patterns of these species.     

Microhabitat use by larvae and females of a rare barrens butterfly,frosted elfin (<Emphasis Type="Italic">Callophrys irus</Emphasis>)

The frosted elfin (Callophrys irus) is a localized and declining butterfly found in xeric open habitats maintained by disturbance. We described the effects of woody plant canopy cover, topography and host plant size and density on the quality of microhabitat of wild indigo (Baptisia tinctoria) host plants containing late instar frosted elfin larvae at four study sites in southeastern Massachusetts, United States. We also assessed whether females preferentially depositing eggs on host plants within specific microhabitats, therefore conferring greater survivorship to the larvae through the late-instar stage. We found that moderate amounts of canopy cover and large plant size characterized larvae-occupied host plants. In the absence of tree canopy cover, late instar larvae density remained low even when host plant density was high. However, females oviposited on wild indigo plants without regard to any of the vegetative or environmental variables we measured. These results indicate that canopy cover was an important characteristic of microhabitats containing late instar larvae, and late instar larvae occupancy was determined by suitable microhabitat conditions, and not female oviposition selection. Managing for canopy cover and microhabitat heterogeneity within relatively open habitats is recommended for the maintenance of frosted elfin populations.     

Predator-induced delayed maturity in bluegill sunfish (Lepomis macrochirus): variation among populations

Previous studies suggested that differences in age at maturity among populations of bluegill sunfish (Lepomis macrochirus) were not genetically based, but rather were a phenotypic response to the presence of predators. I conducted two experiments to determine if the presence of largemouth bass affected age at maturity in bluegill sunfish. Bluegills from three populations were tested to see if the response to the threat of predation varied among source populations. Juvenile bluegills were maintained in the presence of predators or in controls with no contact with predators. Refuge use and growth were monitored during the experiments and reproductive activity was evaluated when bluegills reached age 1. Bluegills from one population exhibited delayed maturity in the presence of predators. Individuals from the other two populations showed no significant differences between predator and control treatments. The population that responded to the presence of predators had a history of high predation levels over the past 30–40 years. The other populations had a history of low levels of predation. This study suggests that presence of predators can induce phenotypic shifts in age at maturity of bluegills, but that the magnitude of response varies among populations in a manner consistent with historical patterns of coexistence. Received: 7 August 1996 / Accepted: 8 August 1997     

阿根廷淡水大米草沼泽中野生豚鼠的种群统计和微生境的利用

本文研究了秋冬季节淡水大米草(Spartina densiflora)沼泽中野生豚鼠(Cavia a perea)的丰度、繁殖和微生境利用,以及其对当地植被和棉鼠类(Sigmodontine)啮齿动物的影响。野生豚鼠喜好S.densiflora覆盖度高的生境。繁殖个体(成体)主要利用矮草为主的斑块,幼体则主要利用禾本科植物为主的斑块。结果说明,野生豚鼠微生境的利用受捕食风险和食物种类的影响。在淡水沼泽中,野生豚鼠的丰度、繁殖、体重和微生境利用没有季节性变化,它们对植物的取食和活动跑道的建造对植被结构和同域的啮齿类动物没有负面影响。通过对具有中度季节性变化的淡水生境中的豚鼠种群和具有高度季节变化的草地和路边中的豚鼠种群进行比较,表明野生豚鼠的种群动态以及豚鼠种群对植被和与其共生的啮齿动物群落的影响都受到冬季植被盖度的限制。