Stage-dependent predation on competitors: consequences for the outcome of a mosquito invasion

1. Predator-mediated coexistence occurs when predation allows competitors to coexist, due to preferential consumption of a superior competitor relative to an inferior competitor. Differences between the native treehole mosquito ( Aedes triseriatus ) and the co-occurring Asian tiger mosquito ( Aedes albopictus ) in anti-predatory larval behaviours account, in part, for the greater vulnerability of this invasive species to native predatory midge ( Corethrella appendiculata ). We test the hypothesis that stage-dependent differences in the sizes of A. albopictus and A. triseriatus larvae, relative to the size-limited C. appendiculata , contribute to differential consumption and the likelihood of predator-mediated coexistence of these competitors.
2. In all instars, larvae of A. triseriatus were larger than A. albopictus of the same stage. Third and fourth instar C. appendiculata selectively consumed late-stage A. albopictus in preference to same-stage A. triseriatus . Small, early-stage prey larvae did not differ in vulnerability to predation, but large, late-stage larvae differed significantly in vulnerability to predation, probably owing to size-limited predation by fourth instar C. appendiculata. This effect was less pronounced for third instar C. appendiculata .
3. Prey size, in conjunction with anti-predatory behavioural responses, alters the probability of predator-mediated coexistence. A stage-structured predation model showed that equally vulnerable early stages reduce the range of environmental conditions (productivities) in which predator-mediated coexistence is possible, increasing the likelihood of both competitive exclusion of the resident species or failure of the invasive to establish. These results underscore the importance of stage-dependent interspecific differences in predator–prey interactions for determining how predators may affect community composition.     

Larval consumption rates,interspecific predation,and local guild composition of egg-overwintering Agabus (Coleoptera,Dytiscidae) species in vernal ponds

Summary In vernal ponds in the boreal region, egg-over-wintering Agabus species form a guild that feeds mainly on larvae and pupae of aedine mosquitoes. The regular co-existence of very similar Agabus species indicates local communities not structured by interspecific competition. However, the lower number of species in local guilds than in the regional species pool poses a problem of limited membership. We suggest that the species of this guild display habitat differences mainly with respect to water temperature, pond size and prey density. In this view, habitat selection reflects body size and thermal growth response of the species, mainly in connection with larval development. We present field data from two northern Swedish vernal ponds. Based on these data, feeding experiments were performed to test the hypothesis outlined above. At a high prey density, larvae of all instars of the larger species A. erichsoni Gemm. & Har. had a significantly higher consumption rate than those of the smaller species A. opacus Aubé. At a low prey density the differences were smaller, and only the third instar larvae differed significantly. At 2° C, larvae of A. opacus had a significantly higher consumption rate than those of A. congener (Thunberg). At 15° C, no significant difference was observed. In studies of within-guild interspecific predation, always the larger larvae consumed the smaller ones. Field data show that egg hatching is spread out in time, and show interspecific differences. Consequently, the effects of unexpected droughts differ with species.     

Field observation of predation on a horsehair worm (Gordioida: Chordodidae) by a diving beetle larva Cybister brevis Aubé (Coleoptera: Dytiscidae)

Diving beetles (Coleoptera: Dytiscidae) are carnivorous in both the larval and adult stages; larvae are exclusively predatory, whereas adults also scavenge for food. They are known to prey on zooplankton, insects, gastropods, fish, amphibians and reptiles. However, there have been no previous reports detailing direct predation on adult Gordioida by Dytiscidae in the field. This study represents the first observation of a diving beetle larva, Cybister brevis Aubé (Coleoptera: Dytiscidae), predating on an adult horsehair worm (Gordioida: Chordodidae). This might be the first report of predation on horsehair worms by insects.     

Pouncing spider, flying mosquito: blood acquisition increases predation risk in mosquitoes

Female mosquitoes dramatically increase their mass when bloodfeeding on their hosts. Such an increase could impact mosquitomortality risk by reducing escape speed and/or agility. We usedtwo laboratory-based experiments to test this notion. In thefirst, we allowed mature female Anopheles gambiae mosquitoesto feed from 0 to 4 min and then attacked those females withan artificial predator. We videotaped subsequent escape responseof each mosquito. Analysis of those responses clearly demonstratedan inverse relationship between increased mass and escape speed.In the second experiment, we exposed both blood-engorged andunfed A.gambiae females to single zebra spiders (Salticus scenicus)in small plexiglass cages. Here, we focused on mosquito escapesfrom searching and pouncing spiders. We found that engorgedmosquitoes were three times less likely to escape searchingspiders compared to unfed conspecifics. Thus we conclude thatblood feeding has substantial state-dependent risk both at thehost (experiment 1) and after feeding (experiment 2). Such riskcan be extended to a broad range of taxa.     

Larval morphology of Agabus clypealis (Thomson, 1867) and A. pseudoclypealis Scholz, 1933 and notes on their distribution (Coleoptera: Dytiscidae)

The third instar larvae of Agabus clypealis (Thomson, 1867) and A. pseudoclypealis Scholz, 1933 are described. Important morphological structures (head, legs, and abdominal segment 8 with urogomphi) are illustrated. Both species share a completely sclerotized abdominal segment 6 with A. setulosus (J. Sahlberg, 1895). Morphological characters and species phenology are discussed in relation to the A. confinis-species group. All available literature records and some new and additional data on distribution of the two species are provided. Agabus clypealis is recorded for the first time from Estonia, and there is no reason to doubt the records of A. pseudoclypealis from Kazakhstan.     

异色瓢虫幼虫对卵的种内自残和种间捕食

自残现象的发生在异色瓢虫Harmonia axyridis中很是常见,为明确自残行为对其存活和生长发育的影响,在非选择性条件下我们对异色瓢虫幼虫对非姊妹卵的种内自残和对七星瓢虫Coccinella septempunctata卵的种间捕食进行了研究。结果表明,异色瓢虫取食同种卵和异种卵均可完成生长发育,与取食蚜虫相比：①其从卵到成虫及蛹期的发育历期明显缩短（P<0.05）,其中1龄幼虫发育历期缩短非常明显（P<0.01）;②自然条件下高死亡率的1龄幼虫的存活率明显提高（P<0.05）;③2-4龄幼虫、蛹和成虫的体重变轻（P<0.05）,且体长在4龄幼虫间也表现出显著差异（P<0.05）。这表明异色瓢虫种内自残和种间捕食对其在食物恶劣条件下延续种群至关重要,但对种群的繁衍却不是最理想的途径。     

Taxonomic review of the genus Copelatus Erichson (Coleoptera: Dytiscidae) in Korea

A taxonomic review of the genus Copelatus Erichson (Coleoptera: Dytiscidae) in Korea is presented. In the present study, five species of Copelatus are recognized from Korea, including one unrecorded species, C. kammuriensis. We provide a key to the Korean species of Copelatus, diagnosis of one unrecorded species, habitus photographs, scanning electron micrographs of the elytron, and illustrations of the aedeagus.     

Selective feeding by larval dytiscids (Coleoptera:Dytiscidae) and effects of fish predation on upper littoral zone macroinvertebrate communities of arctic lakes

Carbon and nitrogen stable isotopic data from the primary producers in mangrove ecosystems are needed to investigate trophic links and biogeochemical cycling. Compared with other mangrove species (e.g. Rhizophora mangle) very few measurements have been conducted on the white mangrove, Laguncularia racemosa. The carbon and nitrogen stable isotopic and elemental compositions of L. racemosa were analyzed and compared from Florida and Belize. ¹³C values of L. racemosa from Florida (mean = –26.4) were slightly higher than those from Twin Cays, Belize (mean = -27.4), which may be due to higher salinity in some parts of the Florida site. There was no difference between the ¹⁵N values from L. racemosa from these two sites (Florida mean = 0.6; Belize mean = 0.3), which are indicative of nitrogen derived from nitrogen fixation in a planktonic marine system. However, higher ¹⁵N values from L. racemosa at Man of War Cay in Belize (11.4 and 12.3), which is fertilized by roosting marine birds (14.0), illustrate that L. racemosa can sensitively reflect alternative nitrogen sources. Although the isotopic data could not distinguish between Avicennia germinans, R. mangle and L. racemosa in Belize the L. racemosa had considerably higher C/N ratios (46.5 – 116.1) compared with the Florida samples (42.2 – 76.0) or the other mangrove species. Unlike some previous findings from R. mangle, substrate characteristics (e.g. salinity, NH₄ ⁺, and H₂S) were not related to the isotopic or elemental composition of L. racemosa. ¹³C, ¹⁵N and C/N were analyzed for ecosystem components from L. racemosa habitats at Twin Cays, including other plants (e.g. R. mangle, A. germinans and seagrass), detritus, microbial mats and sediments. Results from mass-balance calculations show that mangrove detritus composes very little of the sediment, which is principally composed of microbial biomass (80 – 90%). Detritus at some sites is also influenced by sources other than that from L. racemosa, including seagrass leaves.     

First records of Leiodytes nicobaricus (Redtenbacher) and Platambus stygius (Régimbart) in Korea (Coleoptera: Dytiscidae)

Two dytiscid beetle species, Leiodytes nicobaricus (Redtenbacher) and Platambus stygius (Régimbart), are identified for the first time in Korea. Diagnoses, habitus and scanning electron microscopy photographs, and line drawings of the diagnostic characters are provided.     

Selective and frequency dependent predation of aquatic mosquito predator Diplonychus indicus Venkatesan & Rao (Hemiptera: Belostomatidae) on immature stages of three mosquito species

Frequency dependent mosquito larval size (II and IV instars) and species selection by the water bug Diplonychus indicus against three mosquito species Culex quinquefasciatus, Aedes aegypti and Anopheles stephensi was studied in the laboratory. The different frequencies used for each species selection were 20:30:50, 30:50:20, 50:20:30, 25:35:40, 35:40:25 and 40:25:35 of fourth instars of the respective three prey species. All nymphal water bugs (I–V instars) selected IV instar mosquito larvae and the mean proportion of late (larger) larvae eaten by the predator instars was significantly higher than the mean proportion of early (smaller) larvae eaten (F= 2.28; P < 0.001). In all six ratios used to determine the frequency dependent mosquito species selection, all the stages of the water bug selected Ae. aegypti over the other two species (F= 452.43; P < 0.001). The mean number of mosquito larvae eaten increased as its density increased based on various ratios of larvae offered. The study indicated that the predatory efficiency of D. indicus was high when Ae. aegypti was offered as prey, suggesting the utility of this mosquito predator in the control of dengue vectors.     

Molluscan predation by Ovalipes punctatus (De Haan) (Crustacea: Brachyura: Portunidae)

Molluscan predation by the three-spot swimming crab was investigated. The dentition of the heteromorphic chelae allowed crushing, shearing, cutting and holding of prey. Laboratory investigations indicated that small mussels and gastropods were crushed, the larger mussels were prized open, and the foot of the larger gastropods shredded and bits removed. Stomach contents of freshly captured crabs indicated that the crabs are selective carnivores and preferred prey species which are not most abundant in situ (crabs from Kings Beach, Donax serra Röding; crabs from Maitlands River Beach, Bullia rhodostoma Reeve). Ovalipes punctatus (De Haan) foraged on a variety of prey and had no upper prey size limit, but the crabs did show preferences for certain prey sizes. Data indicate that the swimming crabs can effectively utilize the entire mollusc populations on the beaches as prey items.     

Laboratory and field tests of predation on the cereal leaf beetle,Oulema melanopus (Coleoptera: Chrysomelidae)

The cereal leaf beetle (CLB), Oulema melanopus (L.) (Coleoptera: Chrysomelidae), is an invasive pest in North America recently reported in the Canadian Prairies. We performed a series of laboratory assays to identify potential predators and a field study to quantify predation of CLB eggs. In no-choice Petri dish assays, ground beetles (Carabidae), rove beetles (Staphylinidae), and several common lady beetle species (Coccinellidae) were the most consistent predators of eggs and larvae. Nabis spp. (Hemiptera: Nabidae) and wolf spiders (Araneae: Lycosidae) consumed many larvae, but did not consume eggs. Hippodamia spp., Coccinella septempunctata (L.) (Coleoptera: Coccinellidae), and Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) also fed on CLB eggs on potted plants when an alternative food source was available, Sitobion avenae (Fabricius) (Hemiptera: Aphididae). In our field study, we found an average of 24.5% of sentinel eggs disappeared over a 24?h period, likely due to predation. Our results suggest that generalist predators can play an important role in the biological control of CLB, and warrant further study.     

Characterization and localization of Rickettsia sp. in water beetles of genus Deronectes (Coleoptera: Dytiscidae)

In the present study, Rickettsia sp. was detected in four water beetles of the genus Deronectes ( Dytiscidae ) for the first time. Rickettsiae were found in 100% of examined specimens of Deronectes platynotus (45/45), 39.4% of Deronectes aubei (28/71), 40% of Deronectes delarouzei (2/5) and 33.3% of Deronectes semirufus (1/3). Analysis of 16S rRNA gene sequences revealed a phylogenetic relationship with rickettsial isolates of Limonia chorea ( Diptera ), tentatively classified as members of the basal ancestral group. Phylogenetic analysis of the gltA (citrate synthase) gene sequences showed that Deronectes symbionts were closest to bacterial symbionts from spiders. Ultrastructural examinations revealed typical morphological features and intracellular arrangements of rickettsiae. The distribution, transmission and localization of Rickettsia sp. in D. platynotus were studied using a diagnostic PCR assay and FISH. Eggs from infected females of D. platynotus were all Rickettsia -positive, indicative of a vertical transmission.     

Behavior and aquatic plants as factors affecting predation by three species of larval predaceous diving beetles (Coleoptera: Dytiscidae)

Predation among aquatic invertebrate predators can have important effects on patterns of exclusion and coexistence in aquatic habitats, especially if these predators also act as intraguild predators. Such patterns may be explained by variation in predator foraging mode and in the extent and overlap of habitat use. Predaceous diving beetles (Coleoptera: Dytiscidae) are abundant in isolated bodies of water and are effective predators on many aquatic organisms, including other dytiscids. The under-investigated role of hunting behavior and habitat use in altering outcomes of predation under different plant densities may offer insights into patterns of coexistence among larval dytiscids. I performed experiments that quantified behavior of larvae of three common genera of dytiscids that share common prey and then measured predation among genera in the presence or absence of aquatic plants. Behavioral analyses concluded that there were significant differences in foraging modes, with Dytiscus primarily exhibiting sit-and-wait tactics, Graphoderus engaging in active, open water searching, and Rhantus displaying combinations of these behaviors. Predation among larvae was common and occurred when predators were larger than the prey, with no indication of prey preference. Incidence of predation among generic combinations depended on the presence of plants and appeared to be related to behavioral differences among genera. The presence or absence of plants and differences in larval behavior may help to mitigate predation by reducing negative interactions in natural aquatic systems. These results have implications for IGP interactions and may be one of the explanations for the observed richness of this group of predators within aquatic habitats.     

Predation on the invasive mosquito Aedes japonicus (Diptera: Culicidae) by native copepod species in Germany

Some limnic copepod species are predators of mosquito larvae. Seven species belonging to the order Cyclopoida, family Cyclopidae, were collected in the field in Germany and tested for the first time in laboratory bioassays for their potential to serve as biological control agents of the invasive Asian bush mosquito Aedes japonicus (Theobald), a vector of various pathogens causing disease. Females of Diacyclops bicuspidatus (Claus) did not attack 1^st instar larvae of Ae. japonicus, but Macrocyclops distinctus (Richard), Cyclops divergens Lindberg, and C. heberti Einsle predated a mean of 14, 18, or 19 1^st instar larvae, respectively. Acanthocyclops einslei Mirabdullayev and Defaye killed 30 larvae, and high predation rates with a mean of 39 or 46 larvae, respectively, were obtained by Megacyclops viridis (Jurine) and M. gigas (Claus). In regression analyses, predation rates by M. viridis correlated with body size, with specimens of 1.8 mm length being more effective than smaller or bigger ones. Based on the presented data, the two Megacyclops species seem to be promising candidates for use in field studies on the biological control of Ae. japonicus.     

Parasitism of mosquitoes (Diptera: Culicidae) by larval mites (Acari: Parasitengona) in Adelaide, South Australia

Larval mites (Acari) from the cohort Parasitengona were collected from adult female mosquitoes captured in Adelaide, South Australia, from 1997 to 2000. Larvae from three families were identified: Arrenuridae, Hydryphantidae and Erythraeidae. Arrenurid larvae were associated with mosquitoes that use ground pools for larval habitat, while hydryphantids were associated with tree-hole and container-breeding species. Only a single erythraeid record was made. The overall prevalence of parasitism was very low (0.27% of 19 280 mosquitoes) and ranged from 0 to 5.6% for the 16 mosquito species collected. New mite–host records are presented.     

Field observation of foraging behavior by a group of adult diving beetles Agabus (Gaurodytes) bipustulatus preying on an adult Lissotriton boscai

Adult dytiscids prey on a variety of items including other invertebrates but also larger prey such as frogs and fish. Observations of anuran larvae predation are common. However, there have been no reports concerning predation on caudata post‐metamorphosis by dytiscids. Here we describe a predation event by a group of adult diving beetles of the species Agabus (Gaurodytes) bipustulatus (Linnaeus, 1767) on an adult Lissotriton boscai (Lataste in Tourneville, 1879). This report represents the first observation of foraging behavior of adult diving beetles preying on and consuming a living post‐metamorphic newt.     

Sexual dimorphism and sexual conflict in the diving beetle Agabus uliginosus (L.) (Coleoptera: Dytiscidae)

Sexual conflict can drive intersexual arms races, with female resistance and male persistence traits coevolving antagonistically. Such arms races are well documented in some diving beetles, although the extent of sexual conflict in this family remains unclear. The European dytiscid Agabus uliginosus has a strikingly dimorphic female; individuals from most regions are smooth and male‐like, whereas those from some populations have a strongly roughened dorsum, a trait that has attracted the name dispar. We demonstrate that rough and smooth females differ consistently in the development of dorsal surface microreticulation, and that these females are associated with males that differ in the development of their persistence traits. These findings extend the occurrence of pre‐insemination sexual conflict and associated intrasexual dimorphism in Dytiscidae, and suggest that such mating systems are relatively widespread in these beetles.