Clonal mixing in the soldier-producing aphid Pemphigus spyrothecae (Hemiptera: Aphididae)

Illuminating the genetic relationships within soldier-producing aphid colonies is an essential element of any attempt to explain the evolution of the altruistic soldier caste. Pemphigus spyrothecae is a soldier-producing aphid that induces galls on the leaf petioles of its host (trees of the genus Populus). At least a quarter of the aphids within the clonally produced gall population are morphologically and behaviourally distinct first-instar soldiers that defend the gall population from predation. Using field trapping and microsatellites, we investigated the degree of clonal mixing within natural gall populations. Field trapping in the UK showed that all the migrants of P. spyrothecae and of two other Pemphigus species were wingless first-instar soldiers. The average degree of mixing estimated from trapping P. spyrothecae migrants was 0.68% (range = 0-15%). Microsatellite genotyping of 277 aphids from 13 galls collected in Italy revealed an average mixing level of 10.4% (range = 0-59%). Six galls contained more than one clone (range = 2-5 clones). Non-kin aphids were not restricted to the soldier caste but were evenly distributed across instars. An additional gall, from which 527 occupants were genotyped, contained 12 non-kin aphids distributed among nine clones, showing that clonal diversity can be high even when mixing is very low. These observations suggest that although soldiers migrate regularly and can moult and reproduce within foreign galls, clonal mixing in this species is generally low and is unlikely to provide a barrier to the evolution of investment by the aphid clones in an altruistic soldier caste.     

Variation in Escape Behavior of Red and Green Clones of the Pea Aphid

Many insect species have evolved a number of antipredator tactics among which the animal can choose when attacked by a natural enemy. While it is known that individuals may differ in how the antipredator tactics are employed, quantitative studies are rare. In the pea aphid, it has been suggested that different clones differ in their propensity to escape from a predator and that this propensity is linked to the body color of the aphid. We tested clonal variation in the escape behavior in red and green clones of the pea aphid. In three experiments the responses of clones to artificial stimuli and a natural predator were quantified. The results indicate that (1) clones differ considerably in their propensity to show escape behavior, (2) red clones are more likely to drop off the host plant when subjected to an artificial stimulus than green clones are, and (3) the patterns of clonal variation in the escape behavior were not consistent through all three experiments, as clones did not differ in their behavior when attacked by a real predator. The differences in the responses of a clone toward different stimuli supposed to mimic predator attack suggest that extrapolating from laboratory experiments to a field situation may be difficult.     

Social structure and the defensive role of soldiers in a eusocial bamboo aphid,Pseudoregma bambucicola (Homoptera: Aphididae): A test of the defence-optimization hypothesis

A complete understanding of the evolution of sociality in aphids requires a detailed knowledge of the patterns of soldier investment in their ecology. The eusocial bamboo aphidPseudoregma bambucicola has a morphologically specialized first-instar soldier caste. The proportion of soldiers was positively correlated with colony size. Within a colony, soldiers were evenly distributed among subcolonies; within each subcolony, however, their distribution was biased toward peripheries which were exposed to many predators. Field experiments introducing natural enemies such asEupeodes confrater (Diptera: Syrphidae) andSynonycha grandis (Coleoptera: Coccinellidae) revealed that the survival rate of these predators was negatively correlated with the density of soldiers, suggesting that soldiers can more or less defend their colonies by killing or removing a range of natural enemies. Observations suggest that large mature colonies attract more predators than newly established small colonies and that, within a colony, the predators attack each subcolony regardless of its position on bamboo shoots. This implies the presence of a positive correlation between colony size and predation risk. Thus, the investment in soldiers seems to reflect the attacking pattern of predators within a colony. These results agree with the defence-optimization hypothesis in soldier investment ofP. bambucicola colonies.     

Functional response of Chrysoperla carnea (Neuroptera: Chrysopidae) to Helicoverpa armigera (Lepidoptera: Noctuidae): Effect of prey and predator stages

Abstract Understanding predator–prey interactions has a pivotal role in biological control programs. This study evaluated the functional response of three larval instars of the green lacewing, Chrysoperla carnea (Stephens), preying upon eggs and first instar larvae of the cotton bollworm, Helicoverpa armigera Hübner. The first and second instar larvae of C. carnea exhibited type II functional responses against both prey stages. However, the third instar larvae of C. carnea showed a type II functional response to the first instar larvae of H. armigera, but a type III functional response to the eggs. For the first instar larvae of C. carnea, the attack rate on H. armigera eggs was significantly higher than that on the larvae, whereas the attack rate of the second instar C. carnea on H. armigera larvae was significantly higher than that on the eggs. For the third instar larvae of C. carnea, the attack rate on the larvae was 1.015 ± 0.278/h, and the attack coefficient on the eggs was 0.036 ± 0.005. The handling times of the third instar larvae on larvae and eggs were 0.087 ± 0.009 and 0.071 ± 0.001 h, respectively. The highest predation rate was found for the third instar larvae of C. carnea on H. armigera eggs. Results of this study revealed that the larvae of C. carnea, especially the third instar, had a good predation potential in controlling H. armigera eggs and larvae. However, for a comprehensive estimation of the bio‐control abilities of C. carnea toward H. armigera, further field‐based studies are needed.     

Genotypic variation in propensity for host alternation within a population of Pemphigus betae (Homoptera: Aphididae)

In a Utah canyon, the aphid Pemphigus betae shows both a complex life cycle, with alternation between a gall-forming phase on cottonwood leaves and a root-feeding phase on herbs, and a simple life cycle, with year-round residence on roots. In order to determine the extent of clonal variation in life cycle, experiments using multiple sublines of individual clones were carried out in the laboratory and in the field. Previous studies suggested that both genetic and environmental factors underlie life cycle differences among subpopulations of aphids from different sites and different life cycle phases. The current study is the first assessment of clonal variation in propensity for host alternation within a natural population of aphids. In the laboratory experiment, clones showed highly significant differences in reproductive rates and in production of the host-alternating migrants. In agreement with previous findings, clones originating from lineages that had alternated to cotton-wood hosts in the previous year had lower average density and produced more migrants than clones originating from lineages that had remained on roots during the previous year. In order to ascertain how clonal variation and site-specific environmental factors affect life cycle variation under natural conditions, clones from laboratory cultures were used to establish experimental colonies at two elevational sites within the canyon. Production of the host-alternating migrants was affected strongly by clone-x-site interaction and was affected slightly by site. Results from both experiments indicate that loss of host alternation in P. betae could be effected through genetic change, environmental change, or both.     

First record of the adventive oriental aphid Schizaphis piricola (Matsumura, 1917) (Hemiptera, Aphididae) in Europe

The oriental aphid Schizaphis piricola (Matsumura) is recorded for the first time in Europe, on the ornamental pear tree Pyrus calleryana in landscaped areas in Madrid (Spain). Data on the morphology of the forms on primary host (apterous and alate fundatrigeniae and fundatrices), and their biology and distribution are given. The keys for identifying species of Schizaphis (Schizaphis) in the Iberian Peninsula are updated. Two species of aphids are also recorded for the first time on Pyrus calleryana: Schizaphis piricola and Aphis pomi.     

Genetically Distinct Populations in an Asian Soldier-Producing Aphid, Pseudoregma bambucicola (Homoptera: Aphididae), Identified by DNA Fingerprinting and Molecular Phylogenetic Analysis

To estimate genetic structure of a soldier-producing aphid, Pseudoregma bambucicola, samples from natural populations throughout southeastern Asia were analyzed by a DNA fingerprinting technique. We unexpectedly found that P. bambucicola comprises two geographic groups, the northern group and the southern group, which are genetically distinct from each other but morphologically almost indistinguishable. Molecular phylogenetic and statistical analyses based on mitochondrial ribosomal DNA sequences demonstrated that the northern and southern groups of P. bambucicola are not closely related but constitute distinct lineages in the genus Pseudoregma. Detailed morphological reexamination revealed that the two groups could be distinguished by the number of setae on the 8th abdominal tergite of 1st instar nymphs and soldiers. From these results, it was suggested that P. bambucicola should be divided into two species. The northern group from Japan, Taiwan, Hong Kong, and northern Vietnam retains the name P. bambucicola, whereas we suggest that the name P. carolinensis (R. Takahashi, 1941, Tenthredo 3, 208–220) should be used for the southern group from Thailand, Malay Peninsula, Java, Irian Jaya, and Micronesia. The morphological resemblance between P. bambucicola and P. carolinensis might be due to shared ancestral characters of the genus Pseudoregma.     

Spectral composition and visual foraging in the three‐spined stickleback (Gasterosteidae: Gasterosteus aculeatus L.): elucidating the role of ultraviolet wavelengths

Visual signalling can be affected by both the intensity and spectral distribution of environmental light. In shallow aquatic habitats, the spectral range available for visually mediated behaviour, such as foraging, can reach from ultraviolet (UV) to long wavelengths in the human visible range. However, the relative importance of different wavebands in foraging behaviour is generally unknown. Here, we test how the spectral composition of ambient light influences the behaviour of three‐spined sticklebacks (Gasterosteus aculeatus) when foraging for live cladoceran Daphnia magna. Although paying particular attention to the UV waveband, we measured the foraging preferences of sticklebacks for prey presented under four different spectral conditions. These conditions selectively removed UV (UV–), short‐wave (SW–), mid‐wave (MW–) or long‐wave (LW–) light from the entire spectrum. The absence of UV and long wavelengths strongly reduced prey attractiveness for G. aculeatus compared with conditions without short‐wave and mid‐wave light. To control for potential light habitat preferences in the main experiment, we conducted a further choice experiment without prey stimuli. Fish in these trials did not discriminate significantly between the different spectral conditions. When comparing both experiments, it was observed that, although filter preferences for MW– and LW– conditions were virtually consistent, they differed at shorter wavelengths, with a reduced preference for UV– conditions and, at the same time, an increased preference for SW– conditions in the presence of prey. Thus, prey choice seems to be strongly affected by visual information at the short‐wave end of the spectrum. The foraging preferences were also mirrored by the chromatic contrast values between prey and the experimental background, as calculated for each lighting condition using a series of physiological models on stickleback perception. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 359–368.     

The Biology of Tiger Sharks, Galeocerdo Cuvier, in Shark Bay, Western Australia: Sex Ratio, Size Distribution, Diet, and Seasonal Changes in Catch Rates

Tiger sharks, Galeocerdo cuvier, are apex predators in a variety of nearshore ecosystems throughout the world. This study investigates the biology of tiger sharks in the shallow seagrass ecosystem of Shark Bay, Western Australia. Tiger sharks (n = 252) were the most commonly caught species (94%) compared to other large sharks. Tiger sharks ranged from 148–407cm TL. The overall sex ratio was biased towards females (1.8:1), but the sex ratio of mature animals (> 300cm TL) did not differ from 1:1. Contrary to previous accounts, tiger sharks were caught more often in all habitats during daylight hours than at night. Tiger shark catch rates were highly correlated with water temperature and were highest when water temperatures were above 19°C. The seasonal abundance of tiger sharks is correlated to both water temperature and the occurrence of their main prey: sea snakes and dugongs, Dugong dugon. Stomach contents analysis indicated that sea turtles and smaller elasmobranchs were also common prey. The importance of major seagrass grazers (dugongs and green sea turtles, Chelonia mydas) in the diet of tiger sharks suggests the possibility that these sharks are keystone predators in this ecosystem.     

Effect of Manipulations on the Host Selection Behavior of Sitobion avenae (Homoptera: Aphididae)

The study of aphid host selection and feeding behavior is difficult because aphids have to penetrate the plant to reach their feeding site, phloem tissue. The activity of the stylets, salivation or food intake, can not be observed externally and requires an indirect visualization technique such as the Electric Penetration Graph (EPG). The plant selection behavior of Sitobion avenae on potato varied depending on whether an ethological or EPG method was used to study it. A similar variation did not occur with Myzus persicae or Rhopalosiphum padi. The application of water-based silver conductive paint onto the thorax, as normally used for EPG, or onto the abdomen of Sitobion avenae alates resulted in increased duration and frequency of probing compared to results from ethological observations. Our results indicated that EPG manipulations might have different effects on different species of aphids and that a comparison of EPG and ethological data is required to confirm that the EPG method does not bias aphid feeding behavior.     

A new species of Rhopalosiphum (Hemiptera, Aphididae) on Chusquea tomentosa (Poaceae, Bambusoideae) from Costa Rica

The new species Rhopalosiphum chusqueae Pérez Hidalgo & Villalobos Muller, is described from apterous viviparous females caught on Chusquea tomentosa in Cerro de la Muerte (Costa Rica). The identity of the species is supported both by the morphological features and by a molecular phylogenetic analysis based on a fragment of the mitochondrial DNA containing the 5' region of the cytochrome c oxidase 1 (COI) and on the nuclear gene coding for the Elongation factor-1 alpha (EF1α). The taxonomic position of the new species is discussed. An identification key to the Aphidinae species living on plants of Bambusoideae (Poaceae) is presented.     

Stage-Specific Behavioral Responses of Ageneotettix deorum (Orthoptera: Acrididae) in the Presence of Lycosid Spider Predators

Grasshoppers must gather food while avoiding size-selective predation from other arthropods, especially spiders, potentially leading to a trade-off between foraging and defensive behaviors. This trade-off becomes less intense as prey grow larger and are less susceptible to arthropod predation. Activity budgets were constructed for three nymphal (third- to fifth- instar) and adult life cycle stages of Ageneotettix deorum, a common rangeland grasshopper, for three conditions of predation risk by lycosid spiders (spider absence, spider presence, and presence of a nonlethal, chelicerae-modified spider). In third and fourth instars, exposure to predators resulted in reduced feeding activity, increased time spent in antipredator and defensive behaviors, and reduced general activity compared to individuals not exposed to spiders. No significant shifts in behaviors were observed for fifth-instar nymphs and adult A. deorum in response to spider presence. Activity levels in functional spiders and chelicerae-modified spiders were statistically indistinguishable.     

Reproductive isolation among endemic pupfishes (Cyprinodon) on San Salvador Island,Bahamas: microsatellite evidence

Three pupfish (Cyprinodon) morphotypes (two endemic) occur in some of the young (6000 ypb) saline lakes on the Bahamian island of San Salvador. The ‘normal’ morph, a detritivore/omnivore, is not different in its general features from Cyprinodon variegatus from other Bahamian islands. ‘Bulldog’ is a scale‐eater/piscivore that preys upon normal pupfish, and ‘bozo’ is a specialized molluskivore. Reproductive isolation among these morphs is not predicted by the evolutionary biology of congeneric species because sympatry of even morphogically and ecologically quite divergent pupfishes has usually resulted in hybridization/introgression. Survey of variation at eight microsatellite loci reveals that sympatric normal and bulldog populations are genetically distinctive by several criteria, and are therefore likely reproductively isolated. The bulldog morph in Crescent Pond is markedly divergent from those in Little Lake and Osprey Lake, a finding consistent with, although it does not prove, separate parallel origins of this morphotype. The data also suggest that the bulldogs in the latter two lakes did not evolve by intralacustrine speciation from the current sympatric normal populations. Some of the genetic data suggest that the bozo morph may also be reproductively isolated from the other two pupfishes, but only a small, pooled sample of this rare morphotype was available, and the issue is not resolved. Isolating mechanisms between bulldog and normal morphs are of special interest because of the possibility that they arose as a consequence of a predator–prey relationship. A strong correlation between reproductive isolation and predator–prey interactions could provide an important example of ecological speciation via direct selection against heterotypic interactions. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 566–582.     

Effects of prey abundance,distribution, visual contrast and morphology on selection by a pelagic piscivore

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Contact Kairomones Mediating the Foraging Behavior of the Aphid Hyperparasitoid Alloxysta victrix (Westwood) (Hymenoptera: Charipidae)

A variety of host-related substances was assessed in relation to their ability to induce arrestment and antennal examination behavior in adult females of the aphid hyperparasitoid Alloxysta victrix (Westwood). In laboratory bioassays, whole-body homogenates of both unparasitized host aphids [Myzus persicae (Sulzer)] and those parasitized by the primary parasitoid Aphidius colemani Viereck failed to elicit a response. However, significant kairomonal activity was associated with aphid cornicle secretions, aphid honeydew, and solvent extracts of parasitized aphids. Solvent extracts of unparasitized aphids (in either hexane or butan-1-ol) failed to elicit arrestment or antennal examination behavior in hyperparsitoid females. The response to aphid honeydew was significantly lower in experienced females than in naive individuals, suggesting that even relatively strong innate responses can be modified by learning.     

Prey Specificity and the Importance of Close-Range Chemical Cues in Prey Recognition in the Digger Wasp, Liris niger

Females of the digger wasp species Liris niger paralyzed and layed eggs on six of seven tested cricket species. L. niger females with unmanipulated antennae always rejected the cricket Gryllus bimaculatus, but those without antennal flagella paralyzed and layed eggs on it. Even L. niger females from maggots raised artificially on G. bimaculatus never paralyzed it. L. niger first orients toward a prey visually and then touches the prospective prey briefly with its antennal flagella. This contact determines about the suitability of the prey. The ability to differentiate between prey species required at least one partially unmanipulated antennal flagellum. The responsible sensilla were located on the upper surface of the antennal flagella. Chemical cues which make the cricket Acheta domesticus attractive to L. niger females could be transferred to G. bimaculatus.     

Phenotypic plasticity in oysters (Crassostrea virginica) mediated by chemical signals from predators and injured prey

Prey organisms reduce predation risk by altering their behavior, morphology, or life history. Avoiding or deterring predators often incurs costs, such as reductions in growth or fecundity. Prey minimize costs by limiting predator avoidance or deterrence to situations that pose significant risk of injury or death, requiring them to gather information regarding the relative threat potential predators pose. Chemical cues are often used for risk evaluation, and we investigated morphological responses of oysters (Crassostrea virginica) to chemical cues from injured conspecifics, from heterospecifics, and from predatory blue crabs (Callinectes sapidus) reared on different diets. Previous studies found newly settled oysters reacted to crab predators by growing heavier, stronger shells, but that adult oysters did not. We exposed oysters at two size classes (newly settled oyster spat and juveniles ~2.0 cm) to predation risk cue treatments including predator or injured prey exudates and to seawater controls. Since both of the size classes tested can be eaten by blue crabs, we hypothesized that both would react to crab exudates by producing heavier, stronger shells. Oyster spat grew heavier shells that required significantly more force to break, an effective measure against predatory crabs, when exposed to chemical exudates from blue crabs as compared to controls. When exposed to chemical cues from injured conspecifics or from injured clams (Mercenaria mercenaria), a sympatric bivalve, shell mass and force were intermediate between predator treatments and controls, indicating that oysters react to injured prey cues but not as strongly as to cues released by predators. Juvenile oysters of ~ 2.0 cm did not significantly alter their shell morphology in any of the treatments. Thus, newly settled oysters can differentiate between predatory threats and adjust their responses accordingly, with the strongest responses being to exudates released by predators, but oysters of 2.0 cm and larger do not react morphologically to predatory threats.     

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