Factors affecting life and death in Serengeti cheetahs: environment, age, and sociality

We examined environmental and social factors affecting reproductivesuccess across a 20-year data set of individually known cheetahson the Serengeti Plains of Tanzania. Because cheetahs are seeninfrequently and are not amenable to mark–recapture techniques,we devised a model to estimate time of death for individualsthat disappeared from our records. We found that males had markedlylower survival than females. Recruitment was negatively affectedby rainfall but positively affected by numbers of Thomson'sgazelles, the cheetahs' chief prey. There was a negative associationbetween recruitment and numbers of lions, demonstrating thatthe high rates of predation observed in previous studies haveimplications for the dynamics of cheetah populations. Recruitmentwas related to mother's age, peaking when she reached 6–7years. Sociality affected survival in two ways. First, adolescentsliving in temporary sibling groups had higher survival thansingletons, particularly males with sisters. Second, adult malesliving in coalitions had higher survival than singletons inperiods when other coalitions were numerous, yet they had lowersurvival when other coalitions were rare. These results corroborateobservations of enhanced prey capture by female adolescentsand antipredator benefits for adolescents in groups, as wellas competitive advantages for adult males in groups. Furthermore,our findings stress the importance of interactions between environmentaland social factors in affecting reproductive success in mammals.     

Living with the enemy: avoidance of hyenas and lions by cheetahs in the Serengeti

Predator avoidance is likely to play a strong role in structuringspecies communities, even where actual mortality due to predationis low. In such systems, mortality may be low because predatoravoidance is effective, and if the threat of predation is liftedthen entire community structures may be altered. Where competitionis intense, then competitor avoidance may have a similar impacton communities. Avoidance behaviors have been documented fora wide range of species, but this is the first attempt to documentavoidance behavior within a large carnivore community. Audioplayback techniques are used to examine the risk perceivedby cheetahs from their two main competitors that are also theirmain predators, lions and hyenas. The results from these experimentsshow that cheetahs actively moved away from lion and hyenaplayback experiments, compared with dummy playbacks where nosound was played. Cheetahs showed no differences in their responsesto playbacks dependent on their sex or reproductive status,suggesting they were responding principally to a competitionrather than a predation threat. However, cheetahs were muchless likely to hunt after competitor playbacks than after dummyplaybacks, and this resulted in a lower kill rate after competitorplaybacks, demonstrating that the perceived presence of competitorshad a noticeable impact on the foraging rate of cheetahs. Furthermore,while cheetahs moved just as far following lion playbacks asafter hyena playbacks, they spent significantly more time lookingat the loudspeaker and were less likely to make a kill afterlion playbacks, suggesting that cheetahs perceive lions tobe a greater threat than hyenas.     

Where to stay by night and day: Size-specific and seasonal differences in horizontal and vertical distribution of Chaoborus flavicans larvae

1. Data on the distributions of pelagic and benthic Chaoborus flavicans larvae were gathered in 1994 and tested for their agreement with the predator avoidance hypotheses. The development of all Chaoborus life stages, as well as the horizontal and vertical distribution in the four larval instars, was followed from May until October. We expected the largest larvae to dwell deeper by day, thus avoiding predation by visually foraging fish.
2. In agreement with this prediction body size increased with daytime depth, and this was true both between and within instars. The migration amplitude consequently increased with larval instar.
3. There was also evidence for horizontal migration, mainly in the third but also in the fourth instar.
4. Along a horizontal transect with increasing depth, locations with many benthic larvae had fewer pelagic larvae. Oxygen concentration was a good predictor of maximum benthic larval depth for most of the season but failed to predict their distribution in autumn.     

Competition for pulsed resources: an experimental study of establishment and coexistence for an arid-land grass

In arid environments, episodically-pulsed resources are important components of annual water and nutrient supply for plants. This study set out to test whether seedlings have an increased capacity for using pulsed resources, which might then improve establishment when in competition with older individuals. A second aim was to determine whether there is a trade-off in competitive strategies when resources are supplied continuously at low concentrations, or as pulses with pronounced inter-pulse periods. A glasshouse experiment used a target-neighbour design of size-asymmetric competition, with juveniles of Panicum antidotale (blue panicgrass) introduced into contrasting densities of adult plants. Stable isotopes of nitrogen were used for measuring plant resource uptake from pulses, and tolerance to inter-pulse conditions was assessed as the mean residence time (MRT) of nitrogen. A higher root/shoot ratio and finer root system enhanced the capacity of juveniles to use resources when pulsed, rather than when continuously supplied. Higher resource uptake during pulses improved the establishment of juvenile Panicum in mixed cultures with older individuals. However, a trade-off was observed in plant strategies, with juveniles showing a lower MRT for nitrogen, which suggested reduced tolerance to resource deficit during inter-pulse periods. Under field conditions, higher utilization of pulsed resources would lead to the improved seedling establishment of Panicum adjacent to “nurse” plants, whereas mature plants with well-developed roots, exploiting a greater soil volume, maintain more constant resource uptake and retention during inter-pulse periods.     

The change in phototactic behaviour of a Daphnia magna clone in the presence of fish kairomones: the effect of exposure time

Using a clone that responds to the presence of fishkairomones by a pronounced change in phototacticbehaviour, we determined how fast a change to morenegatively phototactic behaviour occurs in Daphnia magnaadults that are exposed to a highconcentration of fish kairomones. Kairomone exposedanimals showed an approximately linear decrease in thevalue of the phototactic index with time. Though theresponse was almost immediate, it took two hoursbefore the difference between fish-induced and controlanimals was significant. Extrapolation of the observedresponse indicates that a maximal change inphototactic behaviour, equivalent to animals that havebeen cultured in the presence of fish kairomones sincebirth, occurs after about 13 hours exposure. Weconclude that the predator-induced change in dielvertical migration of zooplankton is fast, and isfully developed in less than a day. The response timeto fish kairomones of Daphnia is shorter forphototactic behaviour than for life history traits,which may have important consequences with respect tothe evolution of trait-dependence in induced defenceresponses.     

Context-dependent aggregation in Common Frog Rana temporaria tadpoles: influence of developmental stage, predation risk and social environment

1. This study examines the aggregation behaviour and activity of larvae of the Common Frog Rana temporaria in relation to the early social environment, ontogeny and the presence of chemical cues from a predatory fish to address three main questions: (i) Does previous social interaction influence aggregation behaviour in later developmental stages? (ii) To what extent does aggregation behaviour depend upon the risk level perceived by the individual? (iii) Does aggregation behaviour change through ontogeny?
2. Tadpoles were reared from eggs either in small groups or in complete isolation. In relatively early stages of development, tadpoles showed no preference for the side of the test container containing siblings over the side containing no larvae regardless of the social context experienced (isolation or contact with siblings).
3. The presence of chemical cues from a potential predator did not trigger the aggregation behaviour of tadpoles, but it had a remarkable effect on their activity and spatial distribution. Tadpoles exposed to water preconditioned by a predator spent significantly less time swimming and avoided the central area of the test container more frequently than did controls exposed to unconditioned water.
4. Tadpoles were more active, avoided the central section and associated preferentially with conspecifics (siblings) at later stages of development regardless of the social conditioning they had previously experienced.
5. Individuals reared in groups were twice as active as individuals reared in isolation. This suggests that the early social environment experienced by larvae can influence future behaviour and thereby growth and development rates.
6. The expression of conspecific attraction is probably linked to an ontogenetic shift in larval behaviour. However, reduced activity, rather than aggregation, appears to be the basic antipredator mechanism in larval Common Frog.     

Community structure and coexistence of the rotifers of an artificial crater lake

Hydrobiologia - The community structure and dynamics of the rotifers of an abandoned and inundated limestone quarry are described. Zooplankton standing crop is low, but the community is rich in...     

Native species vulnerability to introduced predators: testing an inducible defense and a refuge from predation

To manage the impacts of biological invasions, it is important to determine the mechanisms responsible for the effects invasive species have on native populations. When predation by an invader is the mechanism causing declines in a native population, protecting the native species will involve elucidating the factors that affect native vulnerability. To examine those factors, this study measured how a native species responded to an introduced predator, and whether the native response could result in a refuge from predation. Predation by the green crab, Carcinus maenas, has contributed to the decline in numbers of native soft-shell clams, Mya arenaria, and efforts to eradicate crabs have proven futile. We tested how crab foraging affected clam burrowing, and how depth in the sediment affected clam survival. Clams responded to crab foraging by burrowing deeper in the sediment. Clams at shallow depths were more vulnerable to predation by crabs. Results suggest soft-shell clam burrowing is an inducible defense in response to green crab predation because burrowing deeper results in a potential refuge from predation by crabs. For restoring the native clam populations, tents could exclude crabs and protect clams, but when tents must be removed, exposing the clams to cues from foraging crabs should induce the clams to burrow deeper and decrease vulnerability. In general, by exposing potential native prey to cues from introduced predators, we can test how the natives respond, identify whether the response results in a potential refuge, and evaluate the risks to native species survival in invaded communities.     

Competition between mobile species using patchy resources: an example from a freshwater,symbiotic assemblage

Summary Three species of freshwater mites that are symbiotic with mussels in St Mark's River, north Florida, have consistently high rates of colonization to and occupy high proportions of mussels. The mites Unionicola poundsi and U. serrata are territorial and have limited numbers/host, whereas the non-territorial U. abnormipes has highly variable numbers/host. U. abnormipes and U. serrata are most common in Villosa villosa and Uniomerus declivis respectively, patterns that can not be explained by host species preferences, whereas U. poundsi is equally abundant in both host species. Field experiments showed that both U. poundsi and U. serrata were limited most by intraspecific competition between adult mites, presumably for access to food and oviposition sites. Additionally, U. serrata did not remain within small hosts, most of which were V. villosa. In contrast, numbers of U. abnormipes were limited by both other mite species although the nature of the interactions differed. U. serrata may prey on U. abnormipes when they co-occur, whereas U. poundsi probably only excludes U. abnormipes from certain areas within hosts. Hence, U. abnormipes occurs mostly in V. villosa because most of these mussels do not contain U. serrata, but even so its numbers are still depressed by U. poundsi. The results were consistent with the general expectation of Holmes and Price (1986) that parasite assemblages where species have high colonization levels should be organized primarily by biotic interactions. However, specific outcomes of competition between mites were consistent with the more general model of Levins (1979) for competition between species using variable resources. Failure of other models to apply to Unionicola pinpointed at least five key biological characters that may form a better basis of comparison than taxonomic or habitat-based contrasts.     

Sediments,not plants,offer the preferred refuge for Daphnia against fish predation in Mediterranean shallow lakes: an experimental demonstration

1. Different behavioural responses of planktonic animals to their main predators, fish, have been reported from shallow lakes. In north temperate lakes, large‐bodied zooplankton may seek refuge from predation among macrophytes, whereas in subtropical lakes, avoidance of macrophytes has been observed. The prevalent behaviour probably depends on the characteristics of the fish community, which in Mediterranean lakes is typically dispersed in both the open water zone and in the littoral, as in temperate lakes, and is dominated by small size classes, as in subtropical lakes. 2. We performed ‘habitat choice’ experiments to test the response of Daphnia magna to predation cues at both the horizontal and vertical level by mimicking a ‘shallow littoral’ zone with plants and a ‘deeper pelagic’ zone with sediments. 3. Initial separate response experiments showed that natural plants, artificial plants and predation cues all repelled D. magna in the absence of other stimuli, while sediments alone did not trigger any significant response by D. magna. 4. The habitat choice experiments showed that, in the presence of predation cues and absence of plants, Daphnia moved towards areas with sediment. In the presence of both plants and sediments, Daphnia moved away from the plants towards the sediments under both shallow and deep water treatment conditions. 5. Based on these results, we suggest that Daphnia in Mediterranean shallow lakes avoid submerged macrophytes and instead prefer to hide near the sediment when exposed to predation risk, as also observed in subtropical shallow lakes. This pattern is not likely to change with water level alterations, a common feature of lakes in the region, even if the effectiveness of the refuge may be reduced.     

Competition and stoichiometry: coexistence of two predators on one prey

The competitive exclusion principle (CEP) states that no equilibrium is possible if n species exploit fewer than n resources. This principle does not appear to hold in nature, where high biodiversity is commonly observed, even in seemingly homogenous habitats. Although various mechanisms, such as spatial heterogeneity or chaotic fluctuations, have been proposed to explain this coexistence, none of them invalidates this principle. Here we evaluate whether principles of ecological stoichiometry can contribute to the stable maintenance of biodiverse communities. Stoichiometric analysis recognizes that each organism is a mixture of multiple chemical elements such as carbon (C), nitrogen (N), and phosphorus (P) that are present in various proportions in organisms. We incorporate these principles into a standard predator-prey model to analyze competition between two predators on one autotrophic prey. The model tracks two essential elements, C and P, in each species. We show that a stable equilibrium is possible with two predators on this single prey. At this equilibrium both predators can be limited by the P content of the prey. The analysis suggests that chemical heterogeneity within and among species provides new mechanisms that can support species coexistence and that may be important in maintaining biodiversity.     

Intra- and interspecific density-dependent dispersal in an aquatic prey-predator system

1. Dispersal intensity is a key process for the persistence of prey-predator metacommunities. Consequently, knowledge of the ecological mechanisms of dispersal is fundamental to understanding the dynamics of these communities. Dispersal is often considered to occur at a constant per capita rate; however, some experiments demonstrated that dispersal may be a function of local species density. 2. Here we use aquatic experimental microcosms under controlled conditions to explore intra- and interspecific density-dependent dispersal in two protists, a prey Tetrahymena pyriformis and its predator Dileptus sp. 3. We observed intraspecific density-dependent dispersal for the prey and interspecific density-dependent dispersal for both the prey and the predator. Decreased prey density lead to an increase in predator dispersal, while prey dispersal increased with predator density. 4. Additional experiments suggest that the prey is able to detect its predator through chemical cues and to modify its dispersal behaviour accordingly. 5. Density-dependent dispersal suggests that regional processes depend on local community dynamics. We discuss the potential consequences of density-dependent dispersal on metacommunity dynamics and stability.     

Species coexistence of communities with intraguild predation: The role of refuges used by the resource and the intraguild prey

In this paper, we develop a three-species intraguild predation model which incorporates refuges used by the resource and the intraguild prey, and focus on the effects of refuges on the three species coexistence. The invasion condition and parameter region for coexistence are obtained using invasion analysis. The new invasion condition requires that all boundary states with one missing species can be invaded by the missing species. Numerical simulations show that refuges have a major influence on species coexistence of intraguild predation system, and the results strongly depend on the types of refuges introduced into the model. Our study also shows that prey's refuges are detrimental to species coexistence except the resource using refuges. In contrast to previous research, we find that spatial structure may play an important role in effects of refuges on species coexistence of intraguild predation systems. Our results may shed new light on understanding the mechanisms and the persistence of multi-species predators-prey system.     

Food-web structure and ecosystem services: insights from the Serengeti

The central organizing theme of this paper is to discuss the dynamics of the Serengeti grassland ecosystem from the perspective of recent developments in food-web theory. The seasonal rainfall patterns that characterize the East African climate create an annually oscillating, large-scale, spatial mosaic of feeding opportunities for the larger ungulates in the Serengeti; this in turn creates a significant annual variation in the food available for their predators. At a smaller spatial scale, periodic fires during the dry season create patches of highly nutritious grazing that are eaten in preference to the surrounding older patches of less palatable vegetation. The species interactions between herbivores and plants, and carnivores and herbivores, are hierarchically nested in the Serengeti food web, with the largest bodied consumers on each trophic level having the broadest diets that include species from a large variety of different habitats in the ecosystem. The different major habitats of the Serengeti are also used in a nested fashion; the highly nutritious forage of the short grass plains is available only to the larger migratory species for a few months each year. The longer grass areas, the woodlands and kopjes (large partially wooded rocky islands in the surrounding mosaic of grassland) contain species that are resident throughout the year; these species often have smaller body size and more specialized diets than the migratory species. Only the larger herbivores and carnivores obtain their nutrition from all the different major habitat types in the ecosystem. The net effect of this is to create a nested hierarchy of subchains of energy flow within the larger Serengeti food web; these flows are seasonally forced by rainfall and operate at different rates in different major branches of the web. The nested structure that couples sequential trophic levels together interacts with annual seasonal variation in the fast and slow chains of nutrient flow in a way that is likely to be central to the stability of the whole web. If the Serengeti is to be successfully conserved as a fully functioning ecosystem, then it is essential that the full diversity of natural habitats be maintained within the greater Serengeti ecosystem. The best way to do this is by controlling the external forces that threaten the boundaries of the ecosystem and by balancing the economic services the park provides between local, national and international needs. I conclude by discussing how the ecosystem services provided by the Serengeti are driven by species on different trophic levels. Tourism provides the largest financial revenue to the national economy, but it could be better organized to provide more sustained revenue to the park. Ultimately, ecotourism needs to be developed in ways that take lessons from the structure of the Serengeti food webs, and in ways that provide tangible benefits to people living around the park while also improving the experience of all visitors.     

Sex and the selfish herd: sexual segregation within nonmating whirligig groups

The fitness costs and benefits at different positions in fishshoals, bird flocks, and insect swarms can be asymmetric; agroup's edge may provide more feeding opportunities, but alsogreater predator risk. Animals make trade-offs between theseselection pressures based on individual differences in traitsincluding satiation level, ability to avoid predators, and sex.Previous studies did not evaluate the impact of sex on grouppositioning in these types of nonhierarchical, nonmating groupscalled congregations. A controlled laboratory experiment wasconducted, using marked whirligig beetles (Coleoptera: Gyrinidae),to test for sexual segregation and why different sexes mightchoose different positions. Soon after a disturbance, malesoften were found at the periphery and females at the centerof groups. There was also an overlying influence of feedingon position; satiated individuals moved toward the center andhungry individuals toward the periphery. Several minutes aftera disturbance, sexual segregation disappeared, but segregationdue to hunger persisted. Sexual segregation in this study wasbest explained by the predator avoidance hypothesis, not theenergy needs hypothesis. Females weighed less than males; thismay make them more at risk to predation because of reduced swimmingspeed or less mechanical protection from their exoskeleton.No difference between the sexes was found in the volume of theirdefensive chemicals. This is one of the first studies to showthat sex influences position of individuals within simple nonmatinggroups (congregations) and suggests that more attention shouldbe given to positional sex differences within shoals, flocks,herds, and swarms.     

Temperature- and predator-induced phenotypic plasticity in Bosmina cornuta and B. pellucida (Crustacea: Cladocera)

SUMMARY 1. Clones of Bosmina cornuta and B. pellucida (B. longirostris species complex) were derived from samples collected from Scheuermühlenteich and Lake Windsborn(westernGermany). Experimental temperature change (to 10 °C and 20 °C) and exposure to Acanthocyclops vernalis copepods (12 L^?1) significantly altered external morphology in laboratory cultures of the two species. Morphological traits were derived from eight log₁₀‐transformed and standardised morphometric distances by factorial analysis: factor 1 represented body size, factor 2, size of appendages and factor 3, the head size. 2. Acclimation of clones to cold water (10 °C, >14 days) led to an increase in body, antennule and mucrone size in B. cornuta and B. pellucida. Moreover, at 10 °C, B. cornuta cultures usually collapsed within a few weeks. Compared to the trials at 10 °C, acclimation to 20 °C (the two species) and to 15 °C (B. pellucida only) left the size of body appendages unchanged. Individuals were unequivocally assigned to each species by discriminant functions. Conspecific individuals that were acclimated to different temperatures between 10 and 20 °C also differed in external morphology, but discriminant analysis yielded misclassification rates of 5.3–23.3%. 3. Morphological response to the presence of copepod predators was weaker than that caused by temperature change. Long‐term exposure of clones to copepod predators induced a significant increase in size of appendages in the two species but left body size unaffected. Again, species identification by discriminant functions could be made without any error, whereas conspecific controls and experimentals were misclassified at rates between 19.4 and 29.5%. 4. It is suggested that temperature is the main proximal cue for Bosmina cyclomorphosis. The distinct response to temperature of B. pellucida and B. cornuta may also account for seasonal differences in abundance observed in field.     

Flip,flop and fly: modulated motor control and highly variable movement patterns of autotomized gecko tails

Many animals lose and regenerate appendages, and tail autotomy in lizards is an extremely well-studied example of this. Whereas the energetic, ecological and functional ramifications of tail loss for many lizards have been extensively documented, little is known about the behaviour and neuromuscular control of the autotomized tail. We used electromyography and high-speed video to quantify the motor control and movement patterns of autotomized tails of leopard geckos (Eublepharis macularius). In addition to rhythmic swinging, we show that they exhibit extremely complex movement patterns for up to 30 min following autotomy, including acrobatic flips up to 3 cm in height. Unlike the output of most central pattern generators (CPGs), muscular control of the tail is variable and can be arrhythmic. We suggest that the gecko tail is well suited for studies involving CPGs, given that this spinal preparation is naturally occurring, requires no surgery and exhibits complex modulation.