Tigers eating tigers: evidence of intraguild predation operating in an assemblage of tiger beetles

1. Co‐occurrence of closely related predators in a prey‐limited habitat appears to contradict the principle of competitive exclusion, however it may be explained through indirect effects, niche shifts, and intraguild predation. 2. The interactions between sympatric tiger beetle Cicindela species were examined. Cicindela circumpicta is the largest of three species (C. circumpicta, C. togata, C. fulgida) found in saline habitats throughout central North America. The temporal occurrence of these species overlaps, as does their spatial occurrence on exposed salt flats of saline marshes. During field observations, exoskeletal remains of C. togata were found at the study site in Nebraska, U.S.A. 3. In laboratory trials, male C. circumpicta ate C. togata in 38% of trials and female C. circumpicta ate C. togata in 50% of trials (n = 24). 4. In the field, potential prey, consisting mainly of small flies, was found mostly in shaded conditions but tiger beetles differed significantly in shade use, with C. circumpicta spending 70% of the time in the shade compared with ≈ 20% for C. togata. Differential habitat use was not explained by maximum temperature tolerances, which did not differ between the species. 5. Laboratory trials established that both tiger beetle species consumed small prey (apterous Drosophila) but C. togata was more efficient at capturing winged Drosophila. 6. Foraging efficiency, as measured by the time taken for a C. togata to capture three prey items, decreased significantly in the presence of other tiger beetles, especially C. circumpicta. 7. These results are an indication that intraguild predation and induced changes in foraging behaviour operate in the ecology of adult tiger beetles.     

DNA‐based identification of preys from non‐destructive,total DNA extractions of predators using arthropod universal primers

Here, I show that prey sequences can be detected from DNA of tiger beetles of the genus Rivacindela using whole specimens, nondestructive methods, and universal cytochrome b primers for arthropods. BLAST searches of the obtained sequences against public databases revealed that the diet of Rivacindela is mostly composed of flies but also termites and other beetles. Accurate determination of order, family and even genus was achieved in most cases but rarely to species level. Results suggest that stored DNA samples extracted from whole predatory specimens could be an alternative to dissected gut contents as starting source for DNA‐based dietary studies.     

Alien and native tree species having extrafloral nectaries as favorite hunting area for arboreal endemic Philippine tiger beetles (Coleoptera: Cicindelidae) in human‐disturbed habitat in Lanao del Sur Province,Mindanao, Philippines

To document a relation between abundance of arboreal, predatory tiger beetles, their ant prey, and extrafloral nectaries attracting the ants, we gathered data from more than 10 species of native and introduced trees and large, tree‐like perennial plants in Lanao del Sur Province, Mindanao, Philippines. All specimens of tiger beetles (two Tricondyla and two Neocollyris species, all endemic to the country) were noted on five tree species characterized by presence of extrafloral nectaries, including three alien/invasive and two native ones. Invasive Spathodea campanulata and native Hibiscus tiliaceus were the most inhabited ones (respectively, 56% and 19% of beetles). Presence of tiger beetles on these trees most probably depends on high abundance of ants, which are typical prey for arboreal Cicindelidae, while occurrence of ants can result from presence of extrafloral nectaries on different parts of the plants. This suggests a new mutualistic insect–plant interaction between native and invasive species.     

Feeding Fauna and Foraging Habits of Tiger Beetles Found in Agro‐ecosystems in Western Ghats,India1

Cicindela (Calochroa) whithilli (Hope) and Cicindela (Calochroa) flavomaculata Hope (Cicindelidae: Coleoptera) are seasonally dominant predatory insects in the cultivated, and irrigated rice paddy fields of the South Indian peninsula. While studying the feeding ecology of these tiger beetles in rice paddy agro‐ecosystems in Sringeri area of the central Western Ghats, we examined their potential as biocontrols on the major rice paddy pest populations available in this region. Earthworms and tadpoles were significant prey organisms during the early cultivation period, while other traditional prey organisms like ants and spiders (mainly wolf and jumping spiders) were more common as prey organisms toward the mid‐season. The results showed that the feeding fauna of tiger beetles can extend to prey items beyond arthropods, to include vertebrates. Cicindela (Calochroa) duponti Dejean is another common species that occurs abundantly in the Areca orchards and on the bunds that separate rice paddy fields from adjacent uncultivated lands. We performed laboratory tests to determine the feeding guild of these beetles, and to investigate their possible role as a predator of the rice paddy pests. Although preliminary in nature, the results suggest that both C. whithilli and C. flavomaculata are ineffective as biocontrol agents of rice paddy agro‐ecosystems. They appear to have only a marginal impact on the larvae of Leptocorisa acuta, a major insect pest of paddy rice, but further experimental and observational studies are needed to firmly establish the significance of this observation.     

Prey susceptibilities,prey utilization and variable attack efficiencies of Ural owls

Summary To investigate the factors that influence prey utilization among predators with active prey, three series of experiments were performed in which Ural owls (Strix uralensis) searched for and attacked three prey species of wild mice, Microtus montebelli, Apodemus speciosus, and A. argenteus, in a large flight cage. Over the whole study, owls attacked mice about ten times a night. The number of attacks on each prey species did not differ from that predicted by a random attack model. M. montebelli was taken more than either Apodemus species. Prey utilization appears to be influenced by prey susceptibilities only and it is unlikely that prey selection by the owls affected prey utilization patterns. Under the experimental conditions, random attack is predicted by optimal foraging theory. However, random attack may be explained just as well by the inability of the owl to discriminate prey type. The owls, energy gain was adjusted not by alteration in the number of attacks on a prey species but rather by alteration in the capture success between experiments. Capture success increased in poor food conditions for the same prey species. This flexibility in capture success has not been considered in the assumptions of optimal foraging theory. In conventional optimal foraging theory, the probability of capture success is implicitly assumed as constant and unity. We suggest that this assumption is inadequate to understand the foraging behavior of owls.     

Use of simple remote sensing tools to expedite surveys for rare tiger beetles (Insecta: Coleoptera: Cicindelidae)

Tiger beetles (Insecta: Coleoptera: Cicindelidae) often occupy small patches of suitable habitat in otherwise unsuitable landscapes. Such patches are easily overlooked, which may lead to underestimates of both the number of occurrences and the overall population size. In this study, simple World Wide Web-based tools (Google Earth and Microsoft Terraserver) were used to search high-resolution satellite imagery for patches of suitable habitat for globally and regionally rare tiger beetles on a 3,278 ha wildlife refuge in Maryland, USA. This tract is largely forested but contains scattered small open areas of sand and clay soils that are potential habitat for tiger beetles of conservation concern. Visual inspection of remotely sensed imagery resulted in the identification of 19 potential habitat patches, 15 of which yielded tiger beetle populations when surveyed on the ground. The number of species of tiger beetles recorded from this tract was increased from 3 to 8 and two new sites were discovered for the state sensitive species Cicindela scutellaris rugifrons Dejean. In addition, a small population of C. splendida Hentz was discovered, a species last reported from Maryland in 1948. The technique described here shows great promise for locating small patches of potential tiger beetle habitat in otherwise unsuitable landscapes.     

Species designation of the Bruneau Dune tiger beetle (Cicindela waynei) is supported by phylogenetic analysis of mitochondrial DNA sequence data

Beetles comprise nearly one quarter of described species and show high levels of morphological and ecological diversification. Because of their wide distribution, ease of detection, and correlation of species richness patterns with other taxa, tiger beetles have been recommended for use as a global indicator of regional biodiversity, requiring accurate taxonomic designations. The Bruneau Dune tiger beetle (Cicindela waynei), whose habitat consists of an isolated dune field in southern Idaho, was recently described as a distinct species from the St. Anthony Dunes tiger beetle (C. arenicola) based on morphological characteristics. While these characteristics include distinct differences in genital morphology that could indicate intrinsic reproductive isolation, morphological characteristics have sometimes been misleading in tiger beetle taxonomy. To evaluate genetic support for this species designation, we analyzed 1,751 base pairs of mitochondrial DNA sequence from 147 tiger beetles collected throughout the range of both C. arenicola and C. waynei. Maximum-likelihood and Bayesian phylogenetic analyses indicated monophyly for C. waynei on a well-supported, short branch nested within C. arenicola. Bayesian species delimitation analyses strongly supported C. waynei as a distinct species (speciation probability = 1.0) with the estimated time of divergence ca. 14,500–67,000 years ago. This lack of reciprocal monophyly and recency of speciation is consistent with C. waynei as a member of an evolutionary front where speciation has occurred at a rapid rate. Mitochondrial sequence data supports the species designation of C. waynei, emphasizing the need to determine appropriate management for this species and its restricted habitat.     

Status and conservation of an imperiled tiger beetle fauna in New York State,USA

New York has 22 documented species of tiger beetles (Coleoptera: Cicindelidae). Over half of these species are considered rare, at risk, or potentially extirpated from the state. These rare species specialize on three sandy habitat types under threat from human disturbance: beaches, pine barrens, and riparian cobble bars. In 2005, we began a status assessment of eight of New York’s rarest tiger beetles, examining museum records, searching the literature, and conducting over 130 field surveys of historical and new locations. Significant findings included (1) no detections of four of the eight taxa; (2) no vehicle-free beach habitat suitable for reintroducing Cicindela dorsalis dorsalis; (3) C. hirticollis at only 4 of 26 historical locations; (4) C. patruela patruela at only one site statewide; and (5) C. ancocisconensis at only 3 of 28 de novo survey sites. Additional species that might be declining deserve our attention, as do some threats to tiger beetle habitats, like lack of beach wilderness, fire suppression in pine barrens, and river damming. Rarity in tiger beetles is a result of varying ecologies, which suggest different conservation strategies. Future inventory and documentation of tiger beetle occurrences need to take into account the metapopulation structure and imperfect detectability of these rare insects.     

Intrinsic prey suitability in specialist and generalist Harmonia ladybirds: a test of the trade‐off hypothesis for food specialization

The trade‐off hypothesis posits that increased performance on a given resource comes at the cost of decreased performance on other resources, and that this trade‐off is a driving force of food specialization in both predators and herbivores. In this study, we examined larval survival and performance in two sibling ladybird species, Harmonia yedoensis Takizawa and Harmonia axyridis Pallas (Coleoptera: Coccinellidae), fed on one of four prey species. Harmonia yedoensis is a specialist predator that preys mostly on pine aphids in the field, whereas H. axyridis is a generalist predator with a broad prey range. We experimentally showed in the laboratory that larval survival and performance were not higher when H. yedoensis was fed on pine aphids, compared with the other prey species. Rather, prey suitability was similar in both ladybird species, and H. yedoensis larvae developed as well or even better on prey species that they never utilize in nature. These results suggest that the host range in H. yedoensis may not be limited by the intrinsic suitability of the aphid species per se. Moreover, as shown by our previous study, the pine aphid is a highly elusive prey that is difficult for small ladybird hatchlings to capture, which means that the cost of utilizing this prey is high. Therefore, we conclude that some factor other than prey suitability is responsible for the observed food specialization in H. yedoensis.     

The Effect of Refuge on Dermestes ater (Coleoptera: Dermestidae) Predation on Musca domestica (Diptera: Muscidae): Refuge for Prey or the Predator?

It is believed that habitat heterogeneity can change the extent of predator-prey interactions. Therefore, in this study we examined the effect of habitat heterogeneity (characterized here as an addition of refuge) on D. ater predation on M. domestica. Predation of D. ater on M. domestica larvae was carried out in experimental habitats with and without refuge, and examined at different prey densities. The number of prey eaten by beetles over 24 h of predator-prey interaction was recorded, and we investigated the strength of interaction between prey and predator in both experimental habitats by determining predator functional response. The mean number of prey eaten by beetles in the presence of refuge was significantly higher than in the absence of refuge. Females had greater weight gains than males. Logistic regression analyses revealed the type II functional response for both experimental habitats, even though data did not fit well into the random predator model. Results suggest that the addition of refuge in fact enhanced predation, as prey consumption increased in the presence of refuge. Predators kept in the presence of refuge also consumed more prey at high prey densities. Thus, we concluded that the addition of refuge was an important component mediating D. ater-M. domestica population interactions. Refuge actually acted as a refuge for predators from prey, since prey behaviors detrimental to predators were reduced in this case.     

Spatial variation in the density and vulnerability of preferred prey in the landscape shape patterns of Amur tiger habitat use

Theoretical and empirical research suggests that carnivore distributions are largely determined by prey availability. Availability depends not only on prey density but also on prey accessibility which is affected, in part, by the configuration of landscape attributes that make prey vulnerable to predation. Exactly how spatial variation in these processes shape patterns of carnivore habitat use at the home range scale remains poorly understood. We examined the influence of prey density (negative binomial resource selection function) and vulnerability (kill site resource selection function), mapped separately for each of three species of primary prey, on habitat use patterns within the home range for Amur tigers Panthera tigris altaica in Far East Russia over 20 winters. We developed spatially‐explicit mixed linear regression models to assess these patterns and found that models with parameters for specific primary prey were more robust than models with composite parameters for all primary prey species. This emphasizes the importance of evaluating predation dynamics at a species‐specific level. We also found that Amur tigers used habitat within the home range where red deer Cervus elaphus and wild boar Sus scrofa were dense. These two species were clearly preferred by tigers accounting for 72% (201 of the 278) of the tiger kills detected. The effect of red deer density however, was modulated by the vulnerability of red deer in the landscape. Amur tigers tended to establish their home ranges on habitat where red deer were most vulnerable to predation, but would use habitat where red deer were dense in the peripheral regions of their home ranges. This suggests that tigers may utilize two separate strategies for acquiring prey. As the configuration of resource patches within the home range influences carnivore survival and reproduction, our analysis has implications for tiger conservation that extend beyond our improved understanding of tiger‐prey ecology.     

The feeding behaviour of New Zealand Dolomedes species (Araneae: Pisauridae)

Abstract

The prey spectrum and predatory behaviour of Dolomedes sp. (‘D. III’), D. aquaticus, and D. minor are described from a series of field and laboratory investigations, the former made around Nelson and on Banks Peninsula, Canterbury, during summer months. All species are large, robust spiders that capture prey by directly seizing it in their mouthparts, not using silk at any stage of predation. Adult aquatic insects are their main prey, but these are available only irregularly during their activity period (night-time), and the spiders are opportunistic in their feeding habits. They will eat virtually any available small animal, and at least the largest species, D. III, is able to capture and ingest small fishes. Dead as well as live organisms are taken. Furthermore, the spiders are capable of feeding infrequently; when feeding on small prey organisms they may capture several sequentially, to increase the size of the meal. Live prey is caught while it is in flight, or on the ground, or at the water surface (rarely submerged), and is detected primarily by touch and airborne sound; vision is unnecessary for normal predation. Prey is captured very rapidly, even though this may initially involve a dash of up to 40 cm across the water surface to locate the organism. Stages in the behaviour of an active spider, from waiting for prey to grooming after ingestion, are described.     

A new species of <Emphasis Type="Italic">Laboulbenia</Emphasis> parasitic on tiger beetles

Laboulbenia phaeoxanthae sp. nov. (Ascomycota: Laboulbeniales) parasitic on the tiger beetle Phaeoxantha aequinoctialis bifasciata (Coleoptera: Carabidae: Cicindelinae) from Brazil is described. The new species seems to be closely related to Laboulbenia pheropsophi, but can be distinguished from the latter by the much more slender and elongate habit, and by the different structure of the appendages. The new species is only the second Laboulbeniales species reported on tiger beetles. The widespread Laboulbenia variabilis was previously recorded on Tetracha fulgida from Ecuador.     

The effects of group size on per capita ingestion in flatworms

1. We evaluated the effect of group size on the per capita ingestion rates of three species of flatworm, two of which actively group with conspecifics (Dugesia tigrina, D, dorotocephala) and employ mucus to capture prey, and a third species (Mesostoma ehrenbergii) that does not actively group but does use mucus to capture prey. 2. As flatworm group size increased, daily per capita ingestion first increased and then decreased for D. tigrina and D. dorotocephala. In the case of D. tigrina this pattern was observed even at low predator densities. Ingestion rates of M. ehrenbergii were largely unaffected by group size. 3. Results suggest that the observed changes in per capita ingestion rates with changes in group size previously reported for D. tigrina are related to their tendency for active grouping and are not directly a consequence of prey capture technique or experimental design. 4. We argue that freshwater triclads in general, and D. tigrina in particular, represent an ideal model system for the development and testing of group foraging theory.     

Habitat heterogeneity as the key determinant of the abundance and habitat preference of prey species of tiger in the Chitwan National Park,Nepal

Studies on the relationship between habitat heterogeneity and animal abundance are essential for understanding what determines biodiversity. Transect-based direct observations of eight principal prey species of tiger in the Chitwan National Park (CNP) were used to determine their abundances and habitat preferences. Chital was the most abundant prey species of tiger (Panthera tigris). Each of the prey species had significantly different habitat preferences except sambar deer and chital. Habitat preference was measured using Manly’s preference index, which revealed that short grassland, mixed forest, and riverine forest were the most preferred habitats of the prey species. The results indicate that large species of deer tend to be found in more diverse habitats than small species, except muntjac. The abundance of the principal prey species of tiger was positively correlated with habitat heterogeneity. The habitat, which contributes significantly to the heterogeneity of the landscape, is grassland in large patches of forest. The ongoing increase of forest cover in the CNP has led to a reduction in the area of grassland, which may negatively affect the abundance of the prey species of tiger. Hence, it is suggested that the restoration of landscape heterogeneity is the best way to manage the habitats in the CNP.     

Habitat segregation in four species of adult tiger beetles (Coleoptera: Cicindelidae)

ABSTRACT.

• 1 Habitat segregation in four species of tiger beetles, Cicindela cancellata Dejean, C.cardoni Fleutians, C.minuta Olivier and C.sumatrensis Herbst in a river bank ecosystem was studied in dry and wet seasons.
• 2 The four species segregated distinctly along the river beds into separate habitats, with occasional overlapping in both the seasons.
• 3 Among the habitat characteristics considered, vegetation, soil moisture and available prey-size were found to be important in species segregation.
• 4 D ²-analysis showed that the habitat preferences of C.cardoni and C.cancellata were closer to that of C.sumatrensis. The habitat of C.minuta was distinctly separated.
• 5 There was a positive correlation between the mandible length of each tiger beetle species and the length of prey captured.
• 6 Variance in mandible length within species was related to prey-length distribution pattern and to variance in habitats.

     

Temperature and prey capture: opposite relationships in two predator taxa

Abstract 1. All other things equal, predator capture rates are expected to depend on encounter rate with prey, prey escape capability (including prey defences), and on predator agility. Ectotherm predators and their prey both respond to increasing temperature by increased activity, i.e. predators increase their search area and prey may enhance their escape capability. This means that, as temperature changes, the ability of a predator to catch prey will decrease, increase, or remain unchanged depending on the relative effect of temperature on predator and prey. Their responses may further be differentially moulded by light conditions depending on whether the predator is diurnally or nocturnally active. It was hypothesised that flying Diptera are vulnerable to carabid beetles only at low temperatures and over the full temperature range for spiders because carabids, in contrast to spiders, are not built to catch swiftly moving prey. 2. The first experiment examined the spontaneous locomotor activity of the predators and of fruit flies at different temperatures (5, 10, 15, 20, 25, and 30 °C) and light conditions (light, dark). A second experiment examined the effect of temperature and light on the predation rate of two carabid beetles (Pterostichus versicolor and Calathus fuscipes) and two spiders (Clubiona phragmitis and Pardosa prativaga) using fruit flies (Drosophila melanogaster) as prey. 3. All four predators and the fruit fly increased their locomotory activity at higher temperatures. Activity of the carabid beetles peaked at intermediate temperatures; spiders and fruit flies were most active at the highest temperatures. Predation rate of the spiders increased with temperature whereas the beetles caught flies only at low temperatures (5 and 10 °C). 4. Diurnal variation in temperature may bring different prey groups within the set of potential prey at different times of the day or at different seasons. The ability of many carabid beetles to forage at low temperatures may have nutritional benefits and increases the diversity of interactions in terrestrial food webs.     

Prey preference of large carnivores in Anamalai Tiger Reserve,India

Prey preferences of large carnivores (tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus)) in the tropical forest of Anamalai Tiger Reserve (ATR) were evaluated. This was the first study in ATR to estimate the density of prey and the food habits of these large carnivores. The 958-km² intensive study area was found to have a high mammalian prey density (72.1 animals per square kilometre) with wild boar (20.61 animals per square kilometre) and chital (20.54 animals per square kilometre) being the most common species, followed by nilgiri tahr (13.6 animals per square kilometre). When the density figures were multiplied by the average weight of each prey species, a high biomass density of 14,204 kg km⁻² was obtained for the intensive study area. Scat analysis and incidental kill observation were used to determine the dietary composition of these predators. During the study from the period of March 2001 to April 2004, 1,145 tiger scats, 595 leopard scats and 2,074 dhole scats were collected and analysed. Kill data were based on direct observation of 66 tiger kills and 39 leopard kills. Sambar, with a density of 6.54 kg km⁻² was the preferred prey for these carnivores. Sambar constitutes 35% of the overall diet of tiger, whereas it constitutes 17% and 25% in leopard and dhole diets, respectively. Chital was utilized less than sambar in the range of about 7%, 11% and 15% by tiger, leopard and dhole, respectively. Predator diet was estimated more accurately by scat analysis, which reveals 30% of smaller prey species in leopard’s diet, which was not observed by kill data. This study reveals that ATR harbours high prey density, and these large carnivores seem mostly dependent on the wild prey rather than on domestic livestock as in some other areas in the subcontinent. These factors make ATR a potential area for long-term conservation of these endangered carnivores.