Diving Into The Water: Cues Related to the Decision‐Making by an Egg Parasitoid Attacking Underwater Hosts

Although adult parasitoids spend a majority of their lives above ground, females of several species must search for their host in the water or on the soil. Adult parasitoids above ground can use a variety of sensory cues to detect their hosts from a distance. However, their sensory cues can be impaired from volatile chemicals, and their visual stimuli can be decreased while submerging or burrowing in the water of soil during their search for their hosts. Searching underwater or underground would incur high foraging costs, that is, time and energy consumption and increase risk of drowning. Therefore, to reduce such costs and increase searching efficiency, the decision on where to start submerging or burrowing for attacking hosts is important for parasitoids. Furthermore, there are no studies that have examined the cues of submerging or burrowing parasitoids on their exploit for the decision to attack their hosts. We have examined the cues used by the egg parasitoid Tiphodytes gerriphagus attacking underwater hosts. We compared the searching behaviors of T. gerriphagus among four oviposition site conditions. The four sites investigated were oviposition site with both host adult chemical residues and presence of eggs, with only the presence of eggs, with only the host adult chemical residues, and without any cue. Our results indicated that T. gerriphagus more frequently contacted and submerged at oviposition sites with the adult residues rather than at oviposition sites without them. Nonetheless, the presence of underwater host eggs did not affect the host‐searching behavior. This suggests that T. gerriphagus decided to submerge at the oviposition site in response to the adult residues. Furthermore, our observation also suggested that T. gerriphagus has already detected that the adult residues might be volatile before contacting the oviposition site. Finally, we will discuss the exploitation patterns of host‐searching cue by parasitoids that need to submerge from the context of its reliability and detectability problems.     

Attraction of the sexes inFormica lugubris Zett (Hymenoptera: Formicidae)

Summary Sexuals ofFormica lugubris fly to mating places, where females attract males by using a sex pheromone. Females collected on the nest surface before departing on a mating flight are much less attractive than those collected on the mating place after the mating flight, suggesting that the mating flight triggers the release of the sex pheromone. Olfactory cues are essential for males to locate females while they patrol. Males probably use visual cues to locate females once they have alighted nearby them. Males are also attracted by aggregations of other males on the ground, probably because one or several females are likely to be close to male aggregations.     

Presence of conspecifics triggers host searching behavior in an egg parasitoid wasp

Parasitoids and predators compete for host or prey species. The efficiency of obtaining prey or host items is reduced by intraspecific competition. As the optimal search behavior depends on the intensity of competition, it is important for the parasitoid or predator to obtain information on this intensity. Previous studies have shown that parasitoids can obtain information regarding competition from encounters with already parasitized hosts. They then change their host searching behavior accordingly. However, whether parasitoids obtain the information directly from observing the presence of conspecifics remains unclear. We used Tiphodytes gerriphagus (Marchal) (Hymenoptera: Scelionidae), the solitary egg parasitoid of water striders, for testing the effect of density of conspecifics on host searching behavior. Females of T. gerriphagus dive into the water to search for hosts and sometimes they dive without hosts present. Thus, we investigated whether T. gerriphagus changed underwater activities in response to the density of conspecifics in the absence of hosts. Four densities (1, 2, 4, and 8 female T. gerriphagus) were investigated. Females in competitive situations (groups of 2, 4, and 8 females) displayed host searching behavior, but the solitary females did not. This indicates that the presence of conspecifics triggers host searching behavior and that T. gerriphagus females obtain information on competition directly from conspecifics and use it for modifying their behavior.     

Olfactory host‐finding behaviour of Oulema melanopus (Coleoptera: Chrysomelidae) and its parasitoid,Tetrastichus julis (Hymenoptera: Eulophidae)

Behavioural responses to the host‐associated olfactory cues have not been completely understood for the cereal leaf beetle, Oulema melanopus, and its principal parasitoid, Tetrastichus julis. We, therefore, investigated the role of olfactory cues in the host‐finding behaviour of these species using olfactory bioassays. Behavioural responses of O. melanopus to odours emanating from intact host plants (wheat, oat, barley) vs. a clean‐air control were tested using multichoice and two‐choice bioassays. For T. julis, responses of naïve and experienced adult female wasps to odours associated with the faecal coat of O. melanopus larvae were measured under multichoice and two‐choice conditions. Our results indicate that olfactory cues are involved in the host‐finding behaviour of both O. melanopus and T. julis. Olfactory responses of O. melanopus were influenced by the sex of the beetle and the physiological stage of adults (reproductively active vs. in reproductive diapause). Females respond to olfactory cues in greater proportions than males, and reproductively active, overwintered adults show greater responsiveness than teneral adults in reproductive diapause. Behavioural responses to cues emanating from different crop species were different in multichoice bioassays but not in two‐choice bioassays. Further, we report for the first time that the olfactory cues associated with the faecal coat of O. melanopus evoke host‐finding behaviour of its parasitoid, T. julis. Naïve female wasps are more likely to use these cues to locate the potential host than experienced females. The results of this investigation provide insights into host finding by both the species and the nature of behavioural response brought about by olfactory stimuli, and the results can help to design strategies to improve parasitoid activity by enhancing the crop environment to generate cues for host finding and to manage O. melanopus populations.     

The role of physical cues in the regulation of host recognition and acceptance behavior ofAphidius ervi Haliday (Hymenoptera: Braconidae)

The role of color and shape in the host recognition and acceptance behavior ofAphidius ervi Haliday was studied. A quantitative analysis of the oviposition behavior ofA. ervi was carried out with a computer-aided analysis of 150 video-recorded oviposition sequences on its natural host,Acyrthosiphon pisum (Harris). The importance of visual stimuli was assessed in a choice condition bioassay, observing the behavioral reaction of female parasitoids to various test materials flame-sealed into glass capillaries. Glass beads 2 and 6 mm in diameter and a flat arena were coated with cornicle secretion ofA. pisum, and their acceptance rates by both naive and experienced female parasitoids were assessed under no-choice conditions. In most cases,A. ervi females switched from random searching to attack position when the host was within a range of 1 cm, suggesting that host recognition is regulated in part by cues acting before physical contact. The glass capillary bioassay indicated that visual cues are important factors in the host recognition and acceptance phases. Pea aphid color alone can elicit the oviposition response of naiveA. ervi females, and this response is enhanced when color is combined with aphid shape. The cornicle secretion ofA. pisum stimulated an oviposition response which was stronger in naive females ofA. ervi than in experienced ones and was not significantly affected by the glass bead size or flat surface. These results, along with those from previous studies, suggest that manipulation of the oviposition behavior ofA. ervi is feasible under laboratory conditions.     

Food consumption in Gerris (Hemiptera)

Summary The parameters influencing food consumption in larvae and adults of five species of Gerris were studied. An experimental components analysis was utilized with emphasis placed on measurements of the length of time the insects spent handling food items, and the amount of food in the gut at any one time. The following characteristics were noted: (1) The maximum feeding time required to achieve satiation was 2–2.5 h for all gerrids above 10 mg wet weight. For gerrids below this weight, feeding time declined at a logarithmic rate. (2) Body size had little influence on the duration of the feeding period after food deprivation; only with the first and second instar larvae was a significant difference noted. (3) The average duration of the feeding and non-feeding periods was 10.4 and 24.6 min respectively for satiated adult G. remigis in the presence of excess food. (4) Signifcant differences in relative digestive rate existed between adult and larvae irrespective of species. (5) The smaller the gerrid, the greater the volume relative to its body weight that could be consumed at a single feeding, and for all gerrids, the amount consumed per day was greater than the maximum amount that can be ingested at a single feeding, this difference being larger in the larvae than in the adults. (6) The relationship between daily amount consumed and temperature was linear for four species, increasing with increasing temperature; food consumption in G. remigis, however, peaked at about 19°C, and declined linearily at temperatures both above and below this value. (7) Irrespective of instar and stadium duration, food consumption for G. notabilis peaked within a stadium about 40% of the way through the stadium.     

Developmental times and oviposition rates of predatory mitesTyphlodromus pyri andAmblyseius andersoni (Acari: Phytoseiidae) reared on different foods

The developmental times and the reproduction of two resistant Italian strains ofTyphlodromus pyri Scheuten andAmblyseius andersoni (Chant) were studied in the laboratory by rearing them on the spider mitesPanonychus ulmi (Koch) andEotetranychus carpini (Oud.), on the eriophyidColomerus vitis (Pgst.) and on pollen ofMesembryanthemum criniflorum. The response ofT. pyri andA. andersoni females to a spider mite supply (P. ulmi orE. carpini) of 4, 8 and 16 adult female prey per female predator per day was also studied.Development ofT. pyri onE. carpini andC. vitis required a shorter period than onM. criniflorum pollen, while intermediate values were recorded forP. ulmi. When the highest number of prey was offered, the influence of different foods on oviposition rates ofT. pyri was not significant. An increase in spider mite supply favoured a shorter pre-oviposition period and higher oviposition rates.Development ofA. andersoni was faster on pollen than on spider mites, while intermediate values were found concerningC. vitis. Differences statistically significant were recorded for development onP. ulmi andC. vitis. Colomerus vitis proved to be the more suitable food in terms of oviposition. The oviposition rate decreased when feeding uponP. ulmi, but reached intermediate values onE. carpini andM. criniflorum. Increasing spider mite densities caused shorter pre-oviposition times and higher oviposition rates. Using a given number ofE. carpini females, rather than those ofP. ulmi, resulted in higher oviposition rates and shorter pre-oviposition times.For both predators, the results suggest a higher intrinsic rate of population increase onE. carpini orC. vitis than onP. ulmi.The research was supported by a grant from Regione Veneto (Lotta biologica ed integrata nel controllo di insetti ed acari dannosi).The general lines of the research have been planned by C. Duso. The authors contributed equally to the experimental work.     

Predation in Gerris (Hemiptera): Reactive distances and locomotion rates

Summary Two of the parameters which determine the rate at which prey are encountered by a predator, i.e. the distance at which a predator responds to a prey and its rate of movement relative to the prey's, were determined for all the stages of five species of Gerris using gerrids and Drosophila as prey. These parameters allowed calculation of the swath, or encounter path, a gerrid would cover as it moved across the water surface. Gerris species prefer to attack live prey in front of them, and tend to ignore prey if the attack requires a turn of more than 100°. Hunger was found to affect the responsive angle required to clicit an attack by G. remigis, and regardless of species, smaller gerrids required the prey to be closer before an attack was initiated. The rate of movement in Gerris was measured as a function of stride length and the number of strides made per unit time. Stride length varied according to the length of the mesothoracic leg, and the frequency of movement was observed to be species specific. G. remigis, a stream species, moved 4–6 times as often as the four other species studied, all of which are characteristically found on non-moving water surfaces. Within a species, gerrid size had no significant effect on the frequency of movement, although there was a tendency for smaller gerrids to move less.     

Tetrastylis ovalis: A second case of bat-pollinated passionflower (Passifloraceae)

The floral biology ofTetrastylis ovalis and its pollination by bats was studied in southeastern Brazil. The flowers ofT. ovalis seem more fitted to bat-pollination than those ofPassiflora mucronata, the other known chiropterophilous passionflower. Two species of bats were observed visiting flowers ofT. ovalis: the phyllostomidsGlossophaga soricina andRhinophylla pumilio. Due to the attributes ofTetrastylis, its presumed primitiveness and a better defined bat-pollination syndrome we suggest that chiropterophily evolved independently in the former genus and inPassiflora.     

Role of chemical and visual cues in food recognition by leatherback posthatchlings (Dermochelys coriacea L)

We raised leatherback posthatchlings in the laboratory for up to 7 weeks to study the role of visual and chemical cues in food recognition and food-seeking behavior. Turtles were reared on a formulated (artificial gelatinous) diet and had no contact with test materials until experiments began. Subjects were presented with visual cues (a plastic jellyfish; white plastic shapes [circle, square, diamond] similar in surface area to the plastic model), chemical cues (homogenates of lion's mane jellyfish, Cyanea capillata; moon jellyfish, Aurelia aurita; and a ctenophore, Ocyropsis sp., introduced through a water filter outflow), and visual and chemical cues presented simultaneously. Visual stimuli evoked an increase in swimming activity, biting, diving, and orientation toward the object. Chemical cues elicited an increase in biting, and orientation into water currents (rheotaxis). When chemical and visual stimuli were combined, turtles ignored currents and oriented toward the visual stimuli. We conclude that both cues are used to search for, and locate, food but that visual cues may be of primary importance. We hypothesize that under natural conditions turtles locate food visually, then, as a consequence of feeding, associate chemical with visual cues. Chemical cues then may function alone as a feeding attractant.     

The use of spatial cues for structural guideline orientation inTapinoma sessile andCamponotus pennsylvanicus (Hymenoptera: Formicidae)

Camponotus pennsylvanicus(DeGeer) and Tapinoma sessile(Say) exploit structural elements as guide-lines in their topographic orientation. This research documents the response of T. sessileand C. pennsylvanicusto a series of thigmotactic, gravitational, chemotactic, and phototactic cues while utilizing structural guidelines. Adherence to these guidelines is more pronounced on vertical than on horizontal surfaces and more pronounced in darkness than in daylight. Orientation switches from a crestline on the horizontal to a groove on the vertical. Light and odor trails serve an important role as distance cues in structural guideline orientation. The hierarchy of orientation cues and the adaptive significance of these ant's exploitation of structural guidelines are discussed.     

Lectotypification of Cacao guianensis Aublet (Sterculiaceae)

Hitherto “suppressed,”Cacao guianensis, based on a mixture of elements that can be referred to three currently recognized species (i.e.,Theobroma cacao, T. subincanum, andT. velutinum), is lectotypified by Aublet's illustrations of its flowers. This makesC. guianensis a taxonomic synonym ofT. cacao and encourages nomenclatural stability by permitting the continued use ofT. subincanum andT. velutinum.     

The influence of natural enemies on brood production inIps typographus (Col. scolytidae) with special reference to egg-laying and predation byThanasimus formicarius (Col.: Cleridae)

The predatorThanasimus formicarius (L.) (Coleoptera, Cleridae) and its preyIps typographus (L.) (Coleoptera, Scolytidae) were studied in the laboratory and the field. In the laboratory, 11T. formicarius laid 71–132 eggs (mean=162) during 66–123 days. During this time they ate 66–132I. typographus adults per pair (male +female). The number of eggs laid per female was not correlated with life span or the number ofIps eaten. In the field, predation byT. formicarius larvae and other natural enemies onI. typographus brood was studied in the last year of an outbreak. Caged and uncaged spruce bolts attacked byI. typographus were used, and pairs ofT. formicarius were released in the cages. The treatments were: uncaged bolts, caged bolts withoutT. formicarius, caged bolts with 4T. formicarius pairs, and caged bolts with 8T. formicarius pairs. The productivity ofI. typographus was highest in the caged bolts withoutT. formicarius (mean=4.5 offspring/female) and lowest in the uncaged bolts (mean=0.9 offspring/female). The density ofI. typographus galleries was similar in the different treatments. Hence, the variation in productivity between treatments could not have been due to differences in the levels of intraspecific competition. There was no difference in bark beetle productivity or density ofT. formicarius larvae between bolts with 4 pairs ofT. formicarius and bolts with 8 pairs (mean=2.5 offspring/female). This indicates that some kind of interference occurred betweenT. formicarius individuals (e.g. cannibalism) and that a maximum level of predation was reached. Predation by larvae ofMedetera spp. (Diptera, Dolichopodidae),Thanasimus spp. and other beetles, and parasitism by wasps (Hymenoptera, Pteromalidae) probably caused the low productivity in the uncaged bolts.      

Biology ofTrichogramma ostriniae (Hym.: trichogrammatidae) reared onOstrinia nubilalis (Lep.: pyralidae) and survey for additional hosts

Trichogramma ostriniae has been identified as a candidate for biological control ofOstrinia nubilalis. As little was known about the biology of this parasitoid when reared onO. nubilalis, we undertook experiments to quantify biological parameters important to mass-rearing and ase ofT. ostriniae. When reared continuously onO. nubilalis, femaleT. ostriniae on average lived 2.7 days and produced 24 progeny. Continuous access to honey resulted in a four-fold increase in longevity and fecundity and a significant increase in the percentage of females parasitizing eggs. Rates of fecundity and parasitism decreased with age of female. Likewise, emergence rates and percentage of female progeny decreased with age of parental female.T. ostriniae successfully parasitizedO. nubilalis eggs until the blackhead stage. Most parasitism of eggs and eclosion of adults occurred during the first half of photophase. Eggs of 13 Lepidopterans were parasitized byT. ostriniae. Eggs of the Noctuidae, Pyralidae, and Plutellidae experienced higher levels of parasitism than others tested.T. ostriniae appears to be similar to other species ofTrichogramma in several respects and does not possess any characteristics that limit its potential for mass rearing and use for augmentative biological control ofO. nubilalis.     

Rediscovery of the workerless inquiline antPogonomyrmex colei and additional notes on natural history (Hymenoptera: Formicidae)

Summary Pogonomyrmex colei is a workerless inquiline ant known only from nests ofP. rugosus, its closest relative. Ten of 776 (1.3%) host nests were parasitized at a site in central Arizona, while none of 1499 potential host colonies were parasitized at two other locales. Colonies ofP. colei are perennial, and host alate females in 9 of 10 colonies demonstrates that host queens survive parasitism. Three of 10 colonies died over 19 colony years of observation, while only 1 of 601 colonies became newly parasitized. Mating occurs in morning for up to 2–3 days following summer and fall rains and in afternoon during cool fall days. Mating is intranidal just outside the nest entrance, with males returning to the natal nest. MaleP. colei may be flightless because their wing area is reduced compared to host males. Females fly from the nest and locate potential host colonies by following trunk trails. Workers are the largest barrier to nest establishment, as they removed over 90% ofP. colei females placed in trunk trails or that entered host nests. Males and females ofP. colei andP. anergismus, the only other congeneric inquiline species, are diminutive compared to their hosts, with females 30% lighter than host workers. Fat content is lower and water content is higher inP. colei andP. anergismus females than in their hosts.     

A new species of Tachia (Gentianaceae) from Suriname

A new species ofTachia,T. longipes, is described and illustrated. It is distinct from the other species ofTachia not only in its leaf and stem morphology but also in its characteristic pollen. A revised key to the neotropical species ofTachia is also provided. Studies for theFlora of the Guianas. No. 87.     

Trail-following responses ofTapinoma simrothi (Formicidae: Dolichoderinae) to pygidial gland extracts

Summary The pygidial (anal) gland was found to be the source of trail pheromone in the antTapinoma simrothi. Bioassays conducted with fractionated pygidial gland secretion indicated that the fraction containing iridodials and iridomyrmecin is responsible for the trail pheromone activity. Thus workers ofT. simrothi may utilize the same glandular exudate for alarm and trail following. At high emission rates from a point source, the ants responded in alarm, e.g., rushed to the source with open mandibles and raised abdomen. When concentrations were low and drawn as a line, the ants followed the secretion calmly. Trails ofT. simrothi are long-lived, having a biological half-life of 10 to 19 days. Quantitative studies of the evaporation rates of the iridodials by gas chromatography resulted in a half-life of 11 days, agreeing with the biological data. The implications of the use of the same glandular secretion for alarm and food recruitments are discussed.     

Intraspecific chromosomal polymorphism ofTriticum araraticum (Poaceae) detected by C-banding technique

DifferentTriticum araraticum lines were studied by C-banding method. The intraspecific divergence ofT. araraticum was shown to be caused mainly by large chromosomal rearrangements. Two main chromosomal types were distinguished among the studied lines: (1) a karyotype similar to that ofT. timopheevii and (2) different one. The first type includes some lines ofT. araraticum subspp.kurdistanicum andararaticum; the second comprises most lines ofT. araraticum subsp.araraticum. The lines of the first type can give fertile F₁ hybrids withT. timopheevii.     

Specificity of a strain ofNeozygites sp. (Zygomycetes: Entomophthorales) toMononychellus tanajoa (Acari: Tetranychidae)

Neozygites sp. is commonly found infecting the cassava green mite,Mononychellus tanajoa (Bondar), in parts of northeast Brazil. The introduction of this fungus into other regions requires the knowledge of its specificity, especially in relation to natural enemies of different cassava pests. Laboratory tests indicated the development of germination tubes ofNeozygites in some females ofTetranychus bastosi Tuttle et al. andT. urticae Koch, with subsequent formation of a reduced number of hyphal bodies in someT. bastosi. No females of the phytoseiid predatorsAmblyseius idaeus (Denmark & Muma) andAmblyseius limonicus Garman & McGregor s.l. were infected byNeozygites sp.     

Captive cotton‐top tamarins' (Saguinus oedipus oedipus) use of landmarks to localize hidden food items

Seventeen captive cotton‐top tamarins (Saguinus oedipus oedipus) were individually tested on their use of spatial relationships between landmarks to locate multiple hidden food items. In two experiments, the tamarins were presented with a spatial‐foraging task in which positions of hidden food rewards were fixed in relation to an array of visual cues. In Experiment 1, the cues+hidden food configuration was rotated 90° and the tamarins were successful in locating the food items significantly above chance levels (P<0.01). In Experiment 2 the cues+hidden food configuration was translated (up, down or sideways) from the previously learned configuration, and the monkeys successfully localized the hidden food items (P<0.001). Results indicate that the tamarins relied on the spatial relationship between the multiple landmarks to locate hidden food items rather than on an associative or beacon strategy. The results of these experiments support the contention that when contextually appropriate these captive New World monkeys have the capacity to rely on the spatial relationship or positions of several cues as an array to localize points in their environment. Am. J. Primatol. 71:316–323, 2009. © 2009 Wiley‐Liss, Inc.