Ecological barriers to early colony establishment in three coexisting acacia-ant species in Kenya

In black cotton uplands in East Africa, four symbiotic acacia-ant species compete for possession of a single swollen thorn tree species, Acacia drepanolobium, and yet coexist at fine spatial scales. Three of the four ant species produce independent foundress queens that establish colonies claustrally within swollen thorns, most often on small saplings. We conducted surveys of such saplings at two sites in 2001 and 2004, and examined foundresses and incipient colonies within their swollen thorns to determine what factors influence their success. Competition among foundresses for nest initiation sites was intense, with an average of over one founding attempt per swollen thorn in all samples, and with living and dead queens significantly hyper-dispersed among available thorns. Combat with other foundresses was the most common cause of death among claustral queens, especially for Tetraponera penzigi. In interspecific battles for nest initiation sites, T. penzigi was dominant over Crematogaster nigriceps and C. mimosae, and C. nigriceps won over 80% of its contests with C. mimosae foundresses. For singleton foundresses, brood parasitism by the braconid wasp Trigastrotheca laikipiensis typically results in the death of the entire ant brood. Host queens defend parasite larvae, pupae and eclosed adults, apparently unable to distinguish the wasps from legitimate offspring. Rates of brood parasitism were as high as 15–20% for incipient colonies of both Crematogaster species, but were extremely low for T. nigriceps in all samples. Although T. penzigi and C. nigriceps foundresses are always solitary, approximately 18% of claustral C. mimosae colonies contain cooperating pleometrotic queens. For unparasitized, claustral C. mimosae colonies, brood production per queen did not differ between solitary and cooperating foundresses. However, the per-capita risks associated with parastitism were reduced for pleometrotic queens. Received 8 March 2005; revised 23 May 2005; accepted 3 June 2005.     

Density-dependent effects of multiple predators sharing a common prey in an endophytic habitat

Multiple predator species feeding on a common prey can lead to higher or lower predation than would be expected by simply combining their individual effects. Such emergent multiple predator effects may be especially prevalent if predators share feeding habitat. Despite the prevalence of endophagous insects, no studies have examined how multiple predators sharing an endophytic habitat affect prey or predator reproduction. We investigated density-dependent predation of Thanasimus dubius (Coleoptera: Cleridae) and Platysoma cylindrica (Coleoptera: Histeridae) on a bark beetle prey, Ips pini (Coleoptera: Scolytidae), in a laboratory assay. I. pini utilize aggregation pheromones to group-colonize and reproduce within the stems of conifers. T. dubius and P. cylindrica exploit these aggregation pheromones to arrive simultaneously with the herbivore. Adult T. dubius prey exophytically, while P. cylindrica adults enter and prey within the bark beetle galleries. Larvae of both predators prey endophytically. We used a multiple regression analysis, which avoids confounding predator composition with density, to examine the effects of varying predator densities alone and in combination on herbivore establishment, herbivore reproduction, and predator reproduction. Predators reduced colonization success by both sexes, and decreased I. pini reproduction on a per male and per female basis. The combined effects of these predators did not enhance or reduce prey establishment or reproduction in unexpected manners, and these predators were entirely substitutable. The herbivores net replacement rate was never reduced significantly below one at prey and predator densities emulating field conditions. Similar numbers of each predator species emerged from the logs, but predator reproduction suffered from high intraspecific interference. The net replacement rate of P. cylindrica was not affected by conspecifics or T. dubius. In contrast, the net replacement rate of T. dubius decreased with the presence of conspecifics or P. cylindrica. Combinations of both predators led to an emergent effect, a slightly increased net replacement rate of T. dubius. This may have been due to predation by larval T. dubius on pupal P. cylindrica, as P. cylindrica develops more rapidly than T. dubius within this shared habitat.     

Predation risk of typical ovoid and ‘winged’ morphs of Euplotes (Protozoa,Ciliophora)

Freshwater species of the genus Euplotes (Protozoa, Ciliophora) change their morphology in the presence of some of their predators. The ciliates develop extended lateral wings as well as dorsal and ventral projections which make engulfment by predators more difficult. In a series of laboratory experiments ingestion rates of four protozoan predators, the ciliates Lembadion bullinum, Dileptus anser, Stylonychia mytilus and Urostyla grandis, and one metazoan predator, the turbellarian Stenostomum sphagnetorum, on three species of Euplotes (E. octocarinatus, E. patella and E. aediculatus) were determined. It was calculated that the probability of rejection by a predator changed from 1:1 for ovoid morphs of Euplotes to about 2:1–20:1 for winged morphs of Euplotes, dependent on the prey and predator species that were combined. The nutritional condition of the prey also had some influence. In mixed-species cultures of prey and predators, transformed cells of E. octocarinatus survived for several months.     

Ritualized behavior during competition for food between two formicinae

Summary Polyrhachis laboriosa andCamponotus brutus are two syntopic ants of the African equatorial forest. Although they occupy two different ecological niches (nesting area, diet, rhythms of activity), they are in competition for the exploitation of large permanent food sources. C. brutus, which is nocturnal, changed its rhythm of activity in the presence of large permanent food sources to exploit it day and night, whileP. laboriosa, diurnal, did not change its rhythm of activity. Encounters between workers of the two species at the food source always resulted in duels, even though several other workers were present. When attacked byC. brutus, P. laboriosa workers showed a novel flee-return strategy (fleeing over a 20-cm distance and returning to the source) instead of escaping definitively from the source and displayed ritualized behavior (i.e., fleeing, raising the gaster, flexing the gaster). These types of behavior appeasedC. brutus workers and enabledP. laboriosa to avoid overt aggression and to exploit the source in spite of the presence of a competitor. C. brutus also exhibited ritualized behavior during the duels (i.e., back-and-forth jerking of the body, series of light bites on a leg).C. brutus intimidatedP. laboriosa rather than really attacking it. This ritualization, used at an interspecific level, may be the result of a coevolutionary process or the effect of learning that certain types of behavior are beneficial. In any case, both species benefit from the possibility of exploiting large permanent food sources through confrontations that never lead to overt aggression.     

Laboratory studies on the feeding habits,reproduction and development of three phytoseiid species,Typhlodromus pomi,Phytoseius macropilis andAmblyseius finlandicus (Acari: Phytoseiidae), occurring on abandoned apple trees in Ontario,Canada

Laboratory studies on predation, reproduction and development were conducted on three phytoseiid species,Typhlodromus pomi (Parrott),Phytoseius macropilis (Banks) andAmblyseius finlandicus (Oudemans). These species were dominant phytoseiid mites on the abandoned apple trees with which ecological experiments were carried out in southern Ontario, Canada. Throught the studies, a rust mite species (Diptacus gigantorhynchus sp. complex), a single consistent prey in the orchard, was used as food. Results showed that the three predators developed well with the rust mites as food, and thatA. finlandicus developed most rapidly and had the highest predation rate. However, this species showed a poor tolerance to low prey densities. These observations suggest that this species is a predacious species and the other two species were more or less general feeders. All species showed the same pre-mating behaviour (Amblyseius-Typhlodromus type), but they exhibited a variety of mating and oviposition behaviours.     

Behavior of Adult and Larval Platysoma cylindrica (Coleoptera: Histeridae) and Larval Medetera bistriata (Diptera: Dolichopodidae) During Subcortical Predation of Ips pini (Coleoptera: Scolytidae)

The most abundant predators of the pine engraver, Ips pini (Say), include Platysoma cylindrica (Paykull) and Medetera bistriata Parent. We amended bark sandwiches with pine engravers and adult P. cylindrica and larval M. bistriata, and report observations on these endophytic predators. Adult P. cylindrica preferred adult prey, and appeared to be facultative egg predators, which contrasts with other Histeridae. The larvae of P. cylindrica fed on pine engraver larvae. Protection from predators is generally assumed to be an advantage of an endophytic life history, but we found that this predator exploited the herbivore's gallery architecture to facilitate prey capture. Larval M. bistriata also fed on pine engraver larvae. Immediate paralysis or death of the prey followed a strike with their tentorial rods. Such rapid immobilization, in conjuction with the glandular histology of Medetera spp., suggests that they use a toxin to arrest prey.     

Prey preference of the phytoseiid miteTyphlodromus pyri 1. Response to volatile kairomones

Using a Y-tube olfactometer, a study has been made of the response of females of the predatory miteTyphlodromus pyri Scheuten (Acarina: Phytoseiidae) to volatile kairomones of three prey species: the European red spider mite (Panonychus ulmi (Koch)), the two-spotted spider mite (Tetranychus urticae Koch) and the apple rust mite (Aculus schlechtendali (Nalepa)).Predators that had been reared onT. urticae responded only to the volatile kairomone ofP. ulmi. In contrast, when reared onVicia faba L. pollen, they responded to the kairomones of all three prey species. Pollen-reared predators, offered a choice between kairomones of two different prey species, prefer theP. ulmi kairomone to those ofA. schlechtendali orT. urticae.The difference in response between predators reared onV. faba pollen andT. urticae seems to be caused by the low carotenoid content ofV. faba pollen. Predators that had been reared onV. faba pollen mixed with crystalline -carotene behaved in a way similar to conspecific that had been reared on the carotenoid-rich prey miteT. urticae. Obviously, pollen-rearedT. pyri females are in need of carotenoids, which can be obtained from, e.g.,P. ulmi, T. urticae orA. schlechtendali. This may explain why pollen-reared predators respond to more prey species thanT. urticae-reared predators.WhyT. pyri females need carotenoids has not been established. The only known function of carotenoids in mites is involvement in diapause induction. However, as pollen-rearedT. pyri enter reproductive diapause under short-day conditions, they either extract sufficient amounts of carotenoids fromV. faba pollen, or do not need carotenoids for diapause induction.Apart from the effect of dietary requirements on prey selection, food deprivation also affects the predator's response to kairomones. All the data mentioned above have been obtained for predators that had been starved for 20 h. Predators that had been reared onT. urticae and starved for 48 h before the experiment did respond to the volatile kairomone ofT. urticae in contrast to predators from the same culture that had been starved for 20 h. Thus foraging decisions byT. pyri are affected by both starvation time and specific hunger for carotenoids.     

Feeding behavior ofEuseius stipulatus andTyphlodromus phialatus on the citrus red mitePanonychus citri [Acari: Phytoseiidae,Tetranychidae]

Observations were made on the feeding behavior of the two main phytoseiid species in Spanish Citrus orchards,Euseius stipulatus (Athias-Henriot) andTyphlodromus phialatus Athias-henriot. The experiences were carried out by rearing the predatory mites on excised orange leaves, and always with an excess of the prey the Citrus Red Mite (=CRM)Panonychus citri (McGregor). In experiments with all stages of CRM, the number of prey killed per hour was 5.12 and 2.00, the percentage of successful attacks, 58% and 21%, and the mean time spent feeding on each prey was 5.1 and 12.2 minutes for starving females ofE. stipulatus andT. phialatus respectively.E. stipulatus feeds on all stages of the prey except eggs, andT. philatus, on all stages, except males. Both species attack much less successfully females ofP. citri rather than immatures. In experiments with adult females and eggs ofP. citri as prey, the mean number of prey killed daily was 4.51 females forE. stipulatus, and 2.01 females and 2.12 eggs forT. philatus. Considering this killing rate and the number of eggs laid by the predators in the same period, it can be concluded thatE. stipulatus consumes only 30% of the content of the preys killed, whereasT. philatus consumes a percentage of prey variable between individuals and ranging from 40% to 100%. These differences in feeding behavior between the two species could partly explain differences in their efficiency as biocontrol agents ofP. citri observed in the field.      

Consumption of introduced prey by native predators: Argentine ants and pit-building ant lions

When populations of native predators are subsidized by numerically dominant introduced species, the structure of food webs can be greatly altered. Surprisingly little is known, however, about the general factors that influence whether or not native predators consume introduced species. To learn more about this issue, we examined how native pit-building ant lions (Myrmeleon) are affected by Argentine ant (Linepithema humile) invasions in coastal southern California. Compared to areas without L. humile, invaded areas contained few native ant species and were deficient in medium-sized and large bodied native ants. Based on these differences, we predicted that Argentine ants would negatively affect ant lion larvae. Contrary to this expectation, observational surveys and laboratory growth rate experiments revealed that Myrmeleon were heavier, had longer mandibles, and grew more quickly when their main ant prey were Argentine ants rather than native ants. Moreover, a field transplant experiment indicated that growth rates and pupal weights were not statistically different for larval ant lions reared in invaded areas compared to those reared in uninvaded areas. Argentine ants were also highly susceptible to capture by larval Myrmeleon. The species-level traits that presumably make Argentine ant workers susceptible to capture by larval ant lions—small size and high activity levels—appear to be the same characteristics that make them unsuitable prey for vertebrate predators, such as horned lizards. These results underscore the difficulties in predicting whether or not numerically dominant introduced species serve as prey for native predators.     

Some notes on the ecology of a GermanBranchipus schaefferi population (Crustacea: Anostraca)

Habitat preference, seasonal occurrence, starvation resistance, hatching eggs ofBranchipus schaefferi, and effects of predation onB. schaefferi were studied.Branchipus was only present in turbid, unvegetated ponds and absent in ponds which contain higher aquatic vegetation and theSpirogyra sp. The first individuals ofB. schaefferi appeared in April when water temperature was 10 °C and the last adults in November at a water temperature of 3.5 °C. Up to 6 reproducing generations were observed during this period. Abundance ofB. schaefferi was higher in temporary ponds than in permanent ponds. Sex ratio was close to unity for most of the year. Body size ofB. schaefferi males and females was significantly positively correlated with pond volume. Without foodB. schaefferi could survive for 1.5 to 2 days at 20 °C and 4 to 5 days at 10 °C. Hatching success of eggs decreased when eggs were dried for 7 months. Freezing of eggs had no effect on hatching success. From] the predators tested,Chaoborus sp. larvae clearly selected smallB. schaefferi; one consumed approximately 6Branchipus d–1 at a density of 6 to 12 prey 1–1. The other predators, dragonfly larvae, and larvae and adults ofTriturus alpestris selected alternative prey types, for exampleTubifex sp. and ostracods.     

Different responses of three predatory mite species toTetranychus urticae,Eriophyes dioscoridis andBrevipalpus pulcher: Evidence for the existence of kairomones and allomones

The responses of 3 phytoseiid mite speciesPhytoseiulus persimilis Athias-Henriot,Phytoseius finitimus Ribaga andAmblyseius gossipi Elbadry to allelochemics emitted by prey mite speciesTetranychus urticae Koch,Brevipalpus pulcher (Canestrini & Fanzago) andEriophyes dioscoridis Soliman & Abou-Awad were studied using a test of two-choice assays. The repellent effect elicited by the tenuipalpid mite,B. pulcher and the eriophyid mite,E. dioscoridis againstP. persimilis could be an evidence for the existence of allomones produced by these 2 prey species. The negative response ofP. persimilis to the different stadia ofB. pulcher and the attraction ofP. finitimus toward the same prey suggest that the volatile semiochemicals produced by this prey act as kairomones forP. finitimus and as allomones forP. persimilis. The strong attraction ofP. finitimus andA. gossipi to the different stadia ofT. urticae and the considerable attraction of either predator toB. pulcher compared to the neutral response toE. dioscoridis reveal that both predators show a hierarchy of preference for the kairomones of the 3 prey species studied.      

Aerial prey caching by solitary ground-nesting wasps: A test of the predator defense hypothesis

Many solitary ground-nesting wasps in the families Pompilidae and Sphecidae excavate nests after capturing prey for provisions. These wasps generally cache their immobilized prey temporarily during nest excavation, frequently by suspending the prey in a plant (aerial caching). Here I test the hypothesis that aerial caching by Ammophila spp. wasps (Sphecidae) functions to reduce prey theft by generalist predators, Formica spp. ants. Foraging ants removed baits placed on the ground more rapidly than baits suspended in plants; mean half-lives for ground and aerial baits were 14.5 and 145.7 min, respectively (mean values for experiments 1–3). Ant foraging activity decreased during the midday. Ant interference with nesting activities of Ammophila spp. also decreased during the midday, paralleling observed fluctuations in ant foraging activity.     

Density-dependent predation ofChaoborus flavicans onDaphnia longispina in a small lake: the effect of prey size

Functional response curves of fourth instar larvae ofChaoborus flavicans preying on two size classes ofDaphnia longispina were examined throughout three summer seasons in a small forest lake. Data for each size class were fitted to Holling's disc equation. The parametersa (attack rate) andTh (handling time) were calculated for each prey size from these curves. Attack rate was greater and handling time was shorter for small (0.77 mm) than for large (1.82 mm)Daphnia. In 1:1 mixture of these prey size classes the predation rates ofChaoborus on smallDaphnia at prey densities above 20 l^–1 were greater than predicted from the single size-class experiments. The observed predation rates on largeDaphnia were lower than predicted at all prey densities. Since both single size-class and two size-class experiments were run during the same period of time the difference in observed and predicted predation rates could not be attributed to seasonal changes in prey preference ofChaoborus larvae. In experiments with a concentrated mixture of lake zooplankton (dominated byD. longispina)Chaoborus preference forDaphnia decreased as prey body size increased. There was no obvious correlation between selectivity coefficients and size-frequency distributions ofDaphnia. When medium-sizedDaphnia were omitted from calculations the preference of small over large prey did not differ significantly from the predictions of the single size-class model.     

Searching behavior and associational response ofZetzellia mali (Acarina: Stigmaeidae)

Experimental tests of the hypothesis that natural selection favors predators that search and select prey species that are more profitable are reported. The predator studies was the stigmaeid mite,Zetzellia mali (Ewing). It was observed thatAculus schlechtendali (Nalepa) andPanonychus ulmi (Koch) were more profitable prey thanTetranychus urticae Koch. However,Z. mali attacks prey independently of their nutritional value. Behavioral observations and experiments documented thatZ. mali does not detect kairomones of the prey species. Therefore, prey location is due only to random encounters. Apparently,Z. mali does not search and select prey that are more profitable. The correlation coefficients ofZ. mali, A. schlechtendali, andP. ulmi were investigated in an apple tree. Results revealed that the association depends on season and type of prey.Zetzellia mali andA. schlechtendali were positively associated in the spring (r=0.3,P<0.05) and fall (r=0.36,P<0.01).Zetzellia mali andP. ulmi were not statistically associated. The term associational response is proposed as an ecological process that quantifies the association of predator and prey. The associational-response data suggest thatZ. mali, in the natural ecosystem, responds more to the density ofA. schlechtendali than to the density ofP. ulmi.     

Is prey predation risk influenced more by increasing predator density or predator species richness in stream enclosures?

The direct lethal impacts and the indirect effects predators have on prey characteristics, such as behavior, have fitness consequences for the prey. Whether the level of predation risk that prey face in the presence of multiple predator species can be predicted from a null model that sums the risk from each predator species in isolation is unclear. In field enclosures, we tested whether the predation risk experienced by Stenonema mayfly larvae from a dragonfly larva (Boyeria vinosa) and a hellgrammite (Corydalus cornutus) together matched the predictions of the multiplicative risk model. We then compared whether any deviations from the models predictions were larger in the presence of two predator species than in the presence of an equivalent density of individuals from either predator species alone, to determine if unique effects arise for the prey in the presence of multiple predator species. We also determined if prey moved preferentially into predator-free refuge spaces or decreased their movement in the presence of predators. Stenonemas risk of predation was reduced compared to the models prediction, but no unique multiple predator species effects were present because this risk reduction was comparable in magnitude to the level exhibited in the presence of each predator species alone. The prey did not move into predator-free refuge spaces in the presence of predators in the field enclosures. Thus, these predators appear to interfere interspecifically and intraspecifically, which may facilitate the coexistence of the predators and the prey.     

Systematic implications of the flavonoids and chromosomes ofFlyriella (Compositae—Eupatorieae)

The leaves of five species ofFlyriella were found to contain from one to four glycosides of quercetin and its 4- and 7,4-methyl ethers. These patterns are distinct from those observed for more than seventy species ofBrickellia and support morphological and chromosomal data which indicate thatFlyriella should not be treated as an element ofBrickellia. Alternative treatments are briefly considered.     

The effect of nitrate concentration in a flowing solution system on growth and nitrate uptake of twoPlantago species

Summary Juvenile plants ofPlantago lanceolata andP. major ssp.major were grown in a flowing solution system at 7.5 mM or 9.5 M NO₃. The parameters investigated were: RGR, shoot weight percentage, leaf length, length of main root axis, shoot concentrations of major ions and organic N, and the specific uptake rate for NO₃. At 9.5 M NO₃ growth ofP. major was not hampered, whereas shoot growth and leaf length ofP. lanceolata were reduced. The NO₃ concentration ofP. lanceolata decreased more than that ofP. major. The different performances of the species at 9.5 M NO₃ were associated with different specific uptake rates. In both treatments the root system ofP. major was shorter than that ofP. lanceolata. P. lanceolata accumulated more NO₃ in the leaves. The performance of thePlantago species is discussed in relation to the availability of nutrients in their habitats.Grassland Species Research Group. Publication no. 37.     

Feeding by the aquatic heteropteran,Diplonychus rusticum (Belostomatidae): an effect of prey density on meal size

The behaviour of Diplonychus rusticum feeding on chironomid larvae has been investigated under laboratory conditions. Changes in the percentage of material extracted from prey indicated that feeding for the first two minutes enabled the predator to obtain approximately 33% of the available food; feeding for 10 minutes resulted in only 60% extraction. Comparing the percentage of each prey consumed by D. rusticum exposed to various prey densities, it was apparent that predators were more wasteful and ate less of each prey as chironomid density increased. Because the rate of food intake declined as a greater proportion of each prey was extracted, predators exposed to high chironomid densities reduced the amount of each prey consumed thereby conforming to a simple optimal feeding model.     

Feeding ecology of three predacious aquatic insects and two fish in a riffle of the Speed River,Ontario

The food habits of two fish (Etheostoma flabellare and Rhinichthys atratulus) and three predacious aquatic insects (Nigronia serricornis, Paragnetina media and Agnetina capitata) were examined for one year in a riffle of the Speed River, Ontario. Adult E. flabellare fed primarily on net-spinning caddisflies whereas juvenile E. flabellare relied heavily on chironomids. Rhinichthys atratulus had a more varied diet composed of chironomids, Antocha larvae (Tipulidae) and net-spinning caddisflies. Two of the aquatic insect predators consumed large numbers of net-spinning caddisflies; however, P. media contained large percentages of Hydropsyche and Cheumatopsyche whereas N. serricornis fed primarily on non-hydropsychid net-spinners. The diet of A. capitata was composed largely of chironomids and mayflies. Schoener's diet overlap index () suggested that large E. flabellare and P. media had similar diets; however, E. flabellare ingested much larger prey than P. media. Adult E. flabellare contained a significantly greater proportion of larger prey items in comparison with benthic samples whereas similar comparisons with other predators showed no selection for larger prey. These data demonstrate that E. flabellare was size selective given the range of prey sizes available whereas the other predators were not. Also, there was little dietary overlap among the fish and invertebrate predators suggesting a partitioning of resources.     

Evaluating predation pressure on green treefrog larvae across a habitat gradient

The effect of a predator on the abundance of a prey species depends upon the predators abundance and its ability to capture that prey. The objectives of this research were to evaluate the community structure of predators of green treefrog (Hyla cinerea) tadpoles across habitat types and evaluate the effectiveness of individual predators on H. cinerea tadpoles. Correspondence and cluster analyses of predator frequencies across 23 aquatic habitats indicated that the majority of variance in predator communities was due to a division between permanent and temporary habitats. Experimental work demonstrated that survival of the smallest H. cinerea tadpoles was significantly lower than survival of medium and large tadpoles with the most effective predators, indicating that H. cinerea tadpoles attain a refuge from predation at larger body sizes. We combined the effectiveness of predators in experiments with the abundance of each predator species from the predator community survey to demonstrate that predation pressure on H. cinerea tadpoles is higher in temporary ponds. This pattern may explain in part why this species generally breeds successfully only in permanent habitats. It also confirms that discussions about an increasing gradient of predation pressure from temporary to permanent aquatic habitats should be restricted to individual prey species for which such a gradient has been demonstrated.