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In order to achieve a better understanding of the factors that might have led our hominin ancestors to transition to a more terrestrial niche, including sleeping on the ground, we have conducted a study on the ground nesting behavior of chimpanzees (Pan troglodytes). Chimpanzees, like all other species of great apes, build nests in which to sleep each night, but little is known about regional differences in their nesting habits. Previously, nesting on the ground was considered typical of gorillas, but rare in most populations of chimpanzees. Using data acquired during our extensive chimpanzee nesting survey conducted between 2004 and 2013 across a > 50 000 km2 region in northern Democratic Republic of the Congo, we report a distinctive ground nesting behaviour of eastern chimpanzees (P. t. schweinfurthii). We have mapped the geographical distribution of ground nesting and compared its frequency at 20 survey areas on both sides of a large river, the Uele. We found that ground nests made up more than 1% of total nests at 15 of the 20 survey regions. For a subset of 16 of these regions, we utilized statistical models to investigate whether forest type and structure, as well as the abundance of carnivores and large herbivores, and the activities of humans impacted the frequency of ground nesting and nest height. We predicted that higher encounter rates of human and dangerous animal signs would be associated with lower rates of ground nesting as well as increased nest height. Overall, 10.4% of the Bili-Uéré chimpanzee nests were terrestrial, but the frequency of ground nesting varied extensively between the survey areas (0-29% of nests). The occurrence of ground nests was positively associated with denser forests (p = 0.004), herb patches (p < 0.001), and light gaps (p < 0.001). Light gaps (p < 0.001), herb patches (p = 0.044), and vine tangles (p = 0.016) also had a strong negative effect on nest height. Hunting by humans had a negative effect on the probability of the occurrence of ground nests (p = 0.001) and a positive one on nest height (p = 0.013), with a similar but likely marginal effect of large herbivores on nest height (p = 0.023). In addition, the chimpanzees nested at significantly lower heights with increasing distance from roads and settlements (p < 0.001). Carnivore encounter rates, however, had no significant impact on ground nest frequency or nest height. Our results indicate that ground nesting can no longer be considered a rare and patchily-occurring phenomenon in Pan troglodytes, but is instead a major component of the chimpanzee behavioural repertoire across a considerable fraction of the range of the Eastern subspecies. Our study highlights that neither the large body size of gorillas nor the taming of fire are necessary conditions for hominids to sleep overnight on the ground, even in areas inhabited by multiple species of large carnivore. Human hunting, however, appears to reduce the probability of ground nesting, or eliminate the behavior altogether.

In evolutionarily young species and sympatric host races of phytophagous insects, postzygotic incompatibility is often not yet fully developed, but reduced fitness of hybrids is thought to facilitate further divergence. However, empirical evidence supporting this hypothesis is limited. To assess the role of reduced hybrid fitness, we studied meiosis and fertility in hybrids of two closely related small ermine moths, Yponomeuta padella and Yponomeuta cagnagella, and determined the extent of intrinsic postzygotic reproductive isolation. We found extensive rearrangements between the karyotypes of the two species and irregularities in meiotic chromosome pairing in their hybrids. The fertility of reciprocal F1 and, surprisingly, also of backcrosses with both parental species was not significantly decreased compared with intraspecific offspring. The results indicate that intrinsic postzygotic reproductive isolation between these closely related species is limited. We conclude that the observed chromosomal rearrangements are probably not the result of an accumulation of postzygotic incompatibilities preventing hybridization. Alternative explanations, such as adaptation to new host plants, are discussed.

To improve our understanding of the evolutionary origins of culture and technology in humans, it is vital that we document the full extent of behavioural diversity in our great ape relatives. About half of the world’s remaining chimpanzees (Pan spp.) live in the Democratic Republic of Congo (DRC), yet until now we have known almost nothing about their behaviour. Here we describe the insect-related tool technology of Bili-Uéré chimpanzees (Pan troglodytes schweinfurthii) inhabiting an area of at least a 50,000-km2 area in northern DRC, as well as their percussive technology associated with food processing. Over a 12-year period, we documented chimpanzee tools and artefacts at 20 survey areas and gathered data on dung, feeding remains and sleeping nests. We describe a new chimpanzee tool kit: long probes used to harvest epigaeic driver ants (Dorylus spp.), short probes used to extract ponerine ants and the arboreal nests of stingless bees, wands to dip for D. kohli and stout digging sticks used to access underground meliponine bee nests. Epigaeic Dorylus tools were significantly longer than the other tool types, and D. kohli tools were significantly thinner. Tools classified as terrestrial honey-digging sticks were a significant predictor for brushed and blunted tool ends, consistent with their presumed use. We describe two potential new tool types, an “ant scoop” and a “fruit hammer.” We document an extensive percussive technology used to process termite mounds of Cubitermes sp. and Thoracotermes macrothorax and hard-shelled fruits such as Strychnos, along with evidence of the pounding open of African giant snails and tortoises. We encountered some geographic variation in behaviour: we found honey-digging tools, long driver ant probes and fruit-pounding sites only to the north of the Uele River; there were more epigaeic Dorylus tools to the north and more ponerine ant tools to the south. We found no evidence of termite-fishing, despite the availability of Macrotermes muelleri mounds throughout the region. This lack of evidence is consistent with the results of dung washes, which revealed a substantial proportion of driver ants, but no evidence of Macrotermes or other termites. Our results allow us to describe a new chimpanzee behavioural complex, characterised by a general similarity of multiple behaviours across a large, ecologically diverse region but with subtle differences in prey choice and techniques. We propose that this widespread and related suite of behaviours be referred to as the Bili-Uéré Chimpanzee Behavioural Realm. Possible explanations for this pattern are a recent chimpanzee expansion across the region and the interconnectedness of this population across at least the entirety of northern DRC.

Shelled pteropods are widely regarded as bioindicators for ocean acidification, because their fragile aragonite shells are susceptible to increasing ocean acidity. While short-term incubations have demonstrated that pteropod calcification is negatively impacted by ocean acidification, we know little about net calcification in response to varying ocean conditions in natural populations. Here, we examine in situ calcification of Limacina helicina pteropods collected from the California Current Ecosystem, a coastal upwelling system with strong spatial gradients in ocean carbonate chemistry, dissolved oxygen and temperature. Depth-averaged pH ranged from 8.03 in warmer offshore waters to 7.77 in cold CO2-rich waters nearshore. Based on high-resolution micro-CT technology, we showed that shell thickness declined by ~ 37% along the upwelling gradient from offshore to nearshore water. Dissolution marks covered only ~ 2% of the shell surface area and were not associated with the observed variation in shell thickness. We thus infer that pteropods make thinner shells where upwelling brings more acidified and colder waters to the surface. Probably the thinner shells do not result from enhanced dissolution, but are due to a decline in calcification. Reduced calcification of pteropods is likely to have major ecological and biogeochemical implications for the cycling of calcium carbonate in the oceans.

A key feature of human behavioral diversity is that it can be constrained by cultural preference (“cultural override”); that is, population-specific preferences can override resource availability. Here we investigate whether a similar phenomenon can be found in one of our closest relatives, as well as the potential impacts of ecological differences on feeding behavior. Our study subjects were different subpopulations of Eastern chimpanzees (Pan troglodytes schweinfurthii) occupying two very different habitats, moist tropical lowland forests vs. moist tropical forest–savanna mosaic on opposite sides of a major river. Given differences in encounter rates of different kinds of tool sites on both sides of the Uele River, we predicted that these subpopulations would differ in their likelihood of using tools to prey on two insect species despite similar availability. In surveys conducted over a 9-year period at 19 different survey regions in northern Democratic Republic of Congo (10 in lowland forest and 9 in mosaic), we collected and analyzed data on chimpanzee tool-assisted exploitation of insects. To determine the availability of insect species eaten by the chimpanzees, we counted insects and their mounds on transects and recces at 12 of these sites. For stick tools used to harvest epigaeic Dorylus and ponerine ants, we evaluated seasonal, geographical, and prey-availability factors that might influence their occurrence, using nest encounter rate as a proxy to control for chimpanzee abundance. Across the 19 survey regions spanning both sides of the Uele, we found little difference in the availability of epigaeic Dorylus and ponerine ants. Despite this, tool encounter rates for epigaeic Dorylus, but not ponerine, ants were significantly higher in the mosaic to the north of the Uele. Furthermore, we found no evidence for termite fishing anywhere, despite the availability of Macrotermes mounds throughout the region and the fact that chimpanzees at a number of other study sites use tools to harvest these termites. Instead, the chimpanzees of this region used a novel percussive technique to harvest two other types of termites, Cubitermes sp. and Thoracotermes macrothorax. This mismatch between prey availability and predation is consistent with cultural override, but given the different habitats on the two sides of the Uele River, we cannot fully rule out the influence of ecological factors. Comparing our findings with those of similar studies of other chimpanzee populations promises to contribute to our understanding of the evolution of behavioral diversity in humans and our closest cousins.

The carob moth, Ectomyelois ceratoniae (Zeller; Lepidoptera: Pyralidae), is a devastating pest in high-value crops around the world. An efficient sex pheromone attractant is still missing for the management of this pest, because the major pheromone component is unstable. Host plant volatiles attract herbivore insects and have shown to have good potential to be exploited as alternatives or supplements to sex pheromones. To explore this possibility in carob moth, we assessed the attraction of moths to the volatiles of mature pistachio and different fruit stages of pomegranate, alone and in combination with virgin females, using sticky delta traps in pomegranate orchards of Iran. Traps baited with mature pomegranates, whether uncracked or cracked, infested or uninfested, caught significantly larger numbers of male and both mated and virgin female carob moths than unbaited traps. Traps baited with headspace extract of cracked pomegranate only caught mated females, while mature pistachio only attracted males. Pomegranate flowers, unripe pomegranate, and headspace extract of pistachio did not attract moths. Traps baited with cracked fruit caught more mated females than traps baited with uncracked fruit. Males were attracted similarly to traps baited with cracked-infested pomegranate as to traps baited with virgin females alone. Interestingly, the combination of cracked pomegranate and virgin female enhanced the attraction of virgin females. Together, our results show that volatiles from cracked pomegranates alone or in combination with female sex pheromone have great potential for application in pest management programs of carob moth.

Increasing surface temperatures, Arctic sea-ice loss, and other evidence of anthropogenic global warming (AGW) are acknowledged by every major scientific organization in the world. However, there is a wide gap between this broad scientific consensus and public opinion. Internet blogs have strongly contributed to this consensus gap by fomenting misunderstandings of AGW causes and consequences. Polar bears (Ursus maritimus) have become a "poster species" for AGW, making them a target of those denying AGW evidence. Here, focusing on Arctic sea ice and polar bears, we show that blogs that deny or downplay AGW disregard the overwhelming scientific evidence of Arctic sea-ice loss and polar bear vulnerability. By denying the impacts of AGW on polar bears, bloggers aim to cast doubt on other established ecological consequences of AGW, aggravating the consensus gap. To counter misinformation and reduce this gap, scientists should directly engage the public in the media and blogosphere.

Increasing surface temperatures, Arctic sea-ice loss, and other evidence of anthropogenic global warming (AGW) are acknowledged by every major scientific organization in the world. However, there is a wide gap between this broad scientific consensus and public opinion. Internet blogs have strongly contributed to this consensus gap by fomenting misunderstandings of AGW causes and consequences. Polar bears (Ursus maritimus) have become a “poster species” for AGW, making them a target of those denying AGW evidence. Here, focusing on Arctic sea ice and polar bears, we show that blogs that deny or downplay AGW disregard the overwhelming scientific evidence of Arctic sea-ice loss and polar bear vulnerability. By denying the impacts of AGW on polar bears, bloggers aim to cast doubt on other established ecological consequences of AGW, aggravating the consensus gap. To counter misinformation and reduce this gap, scientists should directly engage the public in the media and blogosphere., BioScience, 68 (4)

Background and Aims The first documented observation of pollination in Pleurothallidinae was that of Endre ́s, who noticed that the ‘viscid sepals’ of Specklinia endotrachys were visited by a ‘small fly’. Chase would later iden- tify the visiting flies as being members of the genus Drosophila. This study documents and describes how species of the S. endotrachys complex are pollinated by different Drosophila species. Methods Specimens of Specklinia and Drosophila were collected in the field in Costa Rica and preserved in the JBL and L herbaria. Flies were photographed, filmed and observed for several days during a 2-year period and were identified by a combination of non-invasive DNA barcoding and anatomical surveys. Tissue samples of the sepals, petals and labellum of Specklinia species were observed and documented by SEM, LM and TEM. Electroantennogram experiments were carried out on Drosophila hydei using the known aggregation pheromones ethyl tiglate, methyl tiglate and isopropyl tiglate. Floral compounds were analysed by gas chromatography–mass spectometry using those same pheromones as standards. Key Results Flowers of S. endotrachys, S. pfavii, S. remotiflora and S. spectabilis are visited and pollinated by several different but closely related Drosophila species. The flies are arrested by aggregation pheromones, including ethyl tiglate, methyl tiglate and isopropyl tiglate, released by the flowers, and to which at least D. hydei is very sen- sitive. Visible nectar drops on the adaxial surface of sepals are secreted by nectar-secreting stomata, encouraging male and female Drosophila to linger on the flowers for several hours at a time. The flies frequently show courtship behaviour, occasionally copulating. Several different Drosophila species can be found on a single Specklinia species. Conclusions Species of the S. endotrachys group share a similar pollination syndrome. There seem to be no spe- cies-specific relationships between the orchids and the flies. It is not expected that Specklinia species will hybridize naturally as their populations do not overlap geographically. The combination of pheromone attraction and nectar feeding is likely to be a generalized pollination syndrome in Pleurothallidinae. Ministerio del Ambiente y Energía/[]/MINAE/Costa Rica Sistema Nacional de Áreas de Conservación/[]/SINAC/Costa Rica Universidad de Costa Rica/[814-B1-239]/UCR/Costa Rica Universidad de Costa Rica/[814-B3-075]/UCR/Costa Rica UCR::Vicerrectoría de Investigación::Unidades de Investigación::Ciencias Agroalimentarias::Jardín Botánico Lankester (JBL)

Several evolutionary explanations for multiple mating in insects have been proposed, and it has been suggested that Lepidoptera benefit from the energy contained in nuptial gifts. In two closely related species of small ermine moths, Yponomeuta cagnagellus (Hübner) and Yponomeuta padellus (L.), we experimentally controlled the number of matings and access to honey as an energy source, and assessed the effects of these factors on female life span and lifetime fecundity. No positive effects of mating were found in either species. In fact, mating significantly shortened life span. The effect of access to energy in the form of honey differed between the two species. In the shorter-lived Yponomeuta padellus, access to honey did not increase the life span of mated females and did not increase fecundity. In Yponomeuta cagnagellus however, access to energy in the form of honey counteracted the negative effects of mating and females with access to honey lived significantly longer and laid significantly more eggs during their lifetime.

In order to assess the impact of human activities on chimpanzee behavior, we compared reactions to humans of Eastern chimpanzees (Pan troglodytes schweinfurthii) living in proximity to and at a distance from roads and settlements in the Bili-Uele landscape in Northern Democratic Republic of the Congo (DRC). We found that chimpanzees in the remote Gangu Forest were more likely to show curious or neutral reactions to us and were less likely to flee than those living closer to roads. In addition, arboreal contact durations with Gangu chimpanzees lasted significantly longer than elsewhere. The most likely explanation for this phenomenon is that with increasing distance from roads, chimpanzees have in the recent past had fewer negative encounters with humans, and thus never learned to fear them. The discovery of this population of naive chimpanzees presents us with an important research and conservation opportunity that may result in the installation of a long-term research site and increased protection of the population.

The degree of polyandry in a species is linked to other life history traits such as egg maturation, life span, and male ejacu- late size and quality. The study of differences in mating strategies between closely related species can provide a better understanding of the evolution of these strategies and of sperm competition. Mating patterns of two closely related species of small ermine moths (Yponomeuta) were investigated in the laboratory. The average female age at first mating was higher in Y. cagnagellus than in Y. padellus. Both species mated more than once; however, Y. cagnagellus females were more likely to remate in a short time frame. Moreover, Y. cagnagellus had higher life time female mating frequencies than Y. padellus (viz., 3.0 versus 2.0). These differences in mating frequency were confirmed in the field by examining the presence of spermatophores (or their remains) in the bursa as well as sperm in the spermatheca of field-caught females.

Anemotaxis in the predatory mite Phytoseiulus persimilis (both well-fed and starved), has previously been studied on a wire grid under slight turbulent airflow conditions yielding weak, yet distinct, gradients in wind speed and odour concentration (Sabelis and Van der Weel 1993). Such conditions might have critically influenced the outcome of the study. We repeated these experiments, under laminar airflow conditions on a flat surface in a wind tunnel, thereby avoiding variation in wind speed and odour concentration. Treatments for starved and well-fed mites were (1) still-air without herbivore-induced plant volatiles (HIPV) (well-fed mites only), (2) an HIPV-free air stream, and (3) an air stream with HIPV (originating from Lima bean plants infested by two-spotted spider mites, Tetranychus urticae). Well-fed mites oriented in random directions in still-air without HIPV. In an air stream, starved mites always oriented upwind, whether plant odours were present or not. Well-fed mites oriented downwind in an HIPV-free air stream, but in random directions in an air stream with HIPV. Only under the last treatment our results differed from those of Sabelis and Van der Weel (1993).

Two compounds present on the surface of Brassica oleracea cv. botrytis leaves have been isolated and identified which stimulate very effectively oviposition in the cabbage root fly, Delia radicum and which are perceived by a specific receptor neuron in the tarsal sensillum C5 of the female fly. Activity of extracts and chromatographic fractions were bioassayed, using oviposition experiments and mainly electrophysiological recordings from the C5 tarsal contact chemoreceptor sensillum of female flies. Spectroscopic data indicate that the main compound is 1,2-dihydro-3-thia-4,10,10b-triaza-cyclopenta[.a.]fluorene-1-carboxylic acid, a novel compound related to Brassica phytoalexins like brassicanal C. It is accompanied by its glycine conjugate.

Summary Small ermine moths (Yponomeutidae: Lepidoptera) are specialist herbivores. Species within the genus Yponomeuta are each specialized on a limited number of plant species, mainly within genera belonging to the Celastraceae. European Yponomeuta species have developed new specialized host affiliations, mainly on rosaceous hosts. Since these host shifts are reputed to be of consequence for speciation, the role of the ovipositing female is of particular interest. Study of the pre-oviposition behaviour of gravid Y. cagnagellus (Hb.) moths on host (Euonymus europaeus), non-host (Crateagus monogyna) and artificial oviposition substrates, provided information on the nature of the cues used for host plant acceptance and the insect’s perception of these cues. Host selection by adult females occurs with contact chemoreceptors probably located on the antennae or tarsi. MeOH-soluble, non-volatile phytochemical compounds washed from the host plant’s surface and applied on an artificial twig are sufficient to stimulate a complete sequence of behavioural elements leading to oviposition. Volatiles do not have a large effect on the pre-oviposition behaviour.

Yponomeuta cagnagellus is a phytophagous moth species specialized on Euonymus europaeus. Host discrimination by the adult female is an important aspect of host specialization and is based mainly on the distinctive secondary chemistry of host and nonhosts. This paper describes a bioassay that was developed to study the effect of isolated plant surface compounds on Yponomeuta oviposition. Adult moths recognize their hosts through chemical stimuli on the leaf or twig surface. Relatively apolar compounds extracted from the host twig surface by washing in dichlormethane do not stimulate oviposition. More polar, methanol-soluble compounds do, and this stimulation is dose dependent. Moths are able to recognize hosts solely by their surface compounds: females show a strong preference for artificial twigs treated with methanolic extracts of their hosts compared to those treated with methanolic extracts of nonhosts Crataegus monogyna and Prunus spinosa (both of which are hosts for closely related Y. padellus). Shape and surface characteristics of the oviposition substrate also influence oviposition. The substrate needs to resemble the basic form of a twig (i.e., cylindrical), and females prefer a coarse surface with irregularities over a smooth one.

The effects of six terpenoids and two terpenoid containing extracts of the neem tree (Azadirachta indica A. Juss.) on oviposition by the diamondback moth (Plutella xylostella L., Yponomeutidae: Lepidoptera) were tested. Two drimane terpenoids, the sesquiterpenoid polygodial and the neem extract Margosan-O exerted significant inhibitory effects at the dosages tested. Ablation experiments showed that both antennae and fore-tarsi contributed to mediation of the inhibition by a drimane. Location of chemosensilla on prothoracic tarsi and ovipositor was examined by scanning electronmicroscopy. Electrophysiological recordings from ovipositor and tarsal taste sensilla showed that distilled water produced distinct responses from one neuron. In tarsal sensilla, ethanol and drimane solutions produced responses from two neurons, one of which might be the water cell that fired at a reduced rate. A drimane significantly decreased the responses of tarsal chemoreceptors to a cabbage leaf extract, which is a possible sensory mechanism leading to behavioural avoidance of this compound.

Contact chemoreception plays a decisive role in host selection and oviposition behaviour of the cabbage root fly, Delia radicum L. (Diptera, Anthomyiidae). Glucosinolates (mustard oil glucosides) are known to be perceived by the flies, and when sprayed on paper leaf-models induce oviposition. Recently it has become clear that other non-volatile types of compounds must also be involved in host selection. A pair of ventro-medial C sensilla on die fifth tarsomere respond strongly to a novel compound called tentatively ‘cabbage identification factor’ (CIF), but not to sucrose, glucose, fructose and proline. CDF is a new non-glucosinolate oviposition stimulant. A single neurone in each sensillum is activated by this compound and the same is true for glucosinolates. In some flies a mixture of bom types of stimuli evoked an apparent mononeural spike train, whereas in odiers spikes of two separate cells were activated. The significance of this variability is not yet clear. The new stimulant, CIF, does not evoke responses in glucosinolate receptors in the D sensilla. The involvement of the C3 sensilla in the detection of host-specific compounds constitutes the first known function for C sensilla in D. radicum. CIF appears to be present in leaf surface extracts from the host-plant Brassica oleracea in quantities as low as 1 ng per gram leaf. In spite of this low level, it stimulates oviposition significantly better than glucobrassicin at higher concentrations, which up till now was known as the most powerful stimulant for D.radicum.

In previous studies we used logistic regression analysis to quantify the change in behavioural phase state of Schistocerca gregaria (Forskal) nymphs subjected to variations in population density. Such work involved restricting insects in small containers either alone or in a crowd. In the present paper we have shown that the fine-scale distribution of food plants, perches and favourable microclimatic sites influences the spatial distribution of locusts, both in the laboratory and under semi-field conditions. When multiple resource sites were provided, solitarious locusts tended to disperse and behavioural gregarization was inhibited. However, provision of only a single site promoted congregation, overcoming the tendency of solitarious insects to avoid each other, and led to behavioural gregarization. The time-course and extent of this response was fully consistent with our earlier experiments using enforced crowding. We suggest that such quantitative, experimental studies of the effects of environmental microstructure on behaviour may yield fundamental insights into the dynamics of plague formation in the desert locust.

The time-course of behavioural change in response to crowding and re-isolation was investigated in adults of Schistocerca gregaria Forskal (Orthoptera, Acrididae) using logistic regression analysis. Crowding solitary-reared adults for a period of 4 h caused them to behave similarly to crowd-reared insects, with their becoming much more active and moving towards rather than away from a stimulus group of locusts. Responsiveness to crowding was greatest in young adults. The behaviour acquired after 48 h of crowding was lost within 1 day of re-isolation. Although experience by solitary-reared adults of crowding for 48 h had only transitory effects on their own behaviour, there was also a long-term influence on the behaviour of their offspring. The strength of this effect was dependent on the age at which adults experienced crowding, increasing in a graded manner with adult age, and hence the recency of crowding before oviposition. Parents crowded at a late stage in the reproductive cycle yielded hatchlings which behaved indistinguishably from those from crowd-reared adults. Such an effect is consistent with the idea that females, through their previous experience of crowding, are effectively predicting the probability that their offspring will emerge into a high-density population, and predisposing their hatchlings' behaviour accordingly.

Pollination based on insect deception has been debated in the scientific community since it was first reported over two hundred years ago. A vast majority of deceptive syndromes occur within the orchid family. While many cheating flowers have been described and are well known, there are still many curious cases that need further investigation. One prime example of such a case is Cypripedium calceolus, known as European lady’s slipper orchid. While the flower has been of interest to many prominent scientists for over a century, its pollination is still not fully understood. Both visual and olfactory cues seem to play an important role in pollinator attraction. In this study we focussed on the olfactory cues in order to explore their relationship (in future experiments) with floral visual cues, including the unique asymmetry of these flowers. Some of the plants’ floral fragrances were used in Electroantennography experiments. Eleven chemical compounds were applied to the antennae of Bombus terrestris and Apis mellifera. Even though these species are not regular visitors of C. calceolus, we were interested to see whether there were common principles in their responses to the flowers’ scent that might justify extrapolating to other pollinator species such as sand bees that get trapped in these orchids and fly out of the flowers afterwards with pollen smeared on their body. The results show that while both species react similarly to most of the odours, some of the tested acetates induced a significantly greater reaction in B. terrestris antennae. These acetates play an important role in bumblebee pheromones, but their relevance for the natural pollinators of C. calceolus remains to be confirmed to see whether chemical mimicry by these flowers is deliberately employed to attract pollinators.

The time-course of behavioural phase change was investigated in nymphs of Schistocerca gregaria, using logistic regression analysis of behaviour recorded in a standard assay. Gregarization occurred very rapidly. Solitary-reared nymphs became markedly gregarious in behaviour within 1-4h of being placed in a crowd. These insects re-solitarized equally quickly if removed from the crowd. Crowd-reared locusts also solitarized within l-4h, but this effect was not complete. Results indicate that, while behavioural gregarization is maximal within a few hours of crowding, solitarization is a two-stage process, changing rapidly at first, then more slowly as a function of the period of previous crowding.

The behavioural phase state of first-instar desert locusts, Schistocerca gregaria Forskal, was quantified by using logistic regression analysis, following treatments in which the population density experienced by their mother during mating and oviposition was manipulated. Crowding during the period of oviposition caused females that had previously been reared in isolation to produce offspring which behaved in a gregarious manner. However, isolating previously crowd-reared females led to hatchlings which behaved more solitariously. The population density experienced during mating affected the behaviour of larvae from solitary-reared, but not crowd-reared, parents. These results show that the transmission of phase characteristics across generations can be modified even at the final stage of the reproductive cycle, providing considerable flexibility in response to changes in population density.

The ultrastructure of a pair of tarsal “C” sensilla located on the ventromedial side near the distal margin of the 5th tarsomere of the female cabbage root fly, Delia radicum (L.) (Diptera : Anthomyiidae) was investigated by electron microscopy and electrophysiological recordings. This “C” sensillum is a typical gustatory sensillum, consisting of a uniporous hair-shaft inserted in a specialized socket and innervated by 5 sensory neurons (i.e. one mechanosensitive and 4 chemosensitive). One of the chemoreceptor cells is sensitive to host-plant compounds, stimulating oviposition. Non-host-plant (carrot) leaf extracts and sucrose did not stimulate any of the receptor cells. Direct contacts between sensory cell somata were observed, and the possibility of peripheral neural interaction at the sensillum level is discussed.

The behavioural phase status of hatchling Schistocerca gregaria Forskal has been quantified using logistic regression, following treatments in which maternal and paternal phase, density during mating and oviposition, and nymphal rearing density were varied. All treatments (having a crowd-reared mother, a crowd-reared father, parents which were crowded only for the period of mating and oviposition, or being reared from birth in a crowd) had a marked gregarizing effect on the behaviour and colour of hatchlings. Such effects have important implications for the generation of locust swarms.

In order to assess the impact of human activities on chimpanzee behavior, we compared reactions to humans of Eastern chimpanzees (Pan troglodytes schweinfurthii) living in proximity to and at a distance from roads and settlements in the Bili-Uele landscape in Northern Democratic Republic of the Congo (DRC). We found that chimpanzees in the remote Gangu Forest were more likely to show curious or neutral reactions to us and were less likely to flee than those living closer to roads. In addition, arboreal contact durations with Gangu chimpanzees lasted significantly longer than elsewhere. The most likely explanation for this phenomenon is that with increasing distance from roads, chimpanzees have in the recent past had fewer negative encounters with humans, and thus never learned to fear them. The discovery of this population of "naïve chimpanzees" presents us with an important research and conservation opportunity that may result in the installation of a long-term research site and increased protection of the population.

Theory predicts that in monandrous butterfly species males should not invest in a long lifespan because receptive females quickly disappear from the mating population. In polyandrous species, however, it pays for males to invest in longevity, which increases the number of mating opportunities and thus reproductive fitness. We tested an extension of this idea and compared male and female lifespan of two closely relatedYponomeutaspecies with different degree of polyandry. Our results confirmed the theoretical prediction that male lifespan is fine-tuned to female receptive lifespan; once-mated males and females of both polyandrous species had an equal lifespan. However, the degree of polyandry was not reflected in male relative to female lifespan. The observed similar female and male lifespan could largely be attributed to a dramatic reduction of female lifespan after mating.

An assay is developed in which the behavioural responses of an individual locust to a group of conspecifics are used to derive an index representing the ‘behavioural phase status’ of that insect. The index is generated from logistic regression models based on 11 behavioural parameters. Data are presented for locusts reared under crowded conditions for many generations and in isolation for one, two and three generations. Crowd-reared locusts differ in behaviour from insects reared in isolation in several ways. They tend to be attracted by a group of locusts, whereas insects reared in isolation are repelled. Solitaryreared locusts also show a characteristic set of behavioural responses which are all consistent with a cryptic lifestyle. Such responses become more evident with increasing number of generations of rearing in isolation. These data represent the most integrated behavioural analysis of locust phase differences yet undertaken. The procedure will be used in future work to quantify the dynamics of gregarization in congregated solitarious locusts. This will provide a much-needed basis for studying underlying controlling mechanisms, the nature of which is at present unclear.

The role of glucosinolates in the oviposition behaviour of the cabbage root fly, Delia radicum (L.) (Diptera, Anthomyiidae) was investigated using egg counts and electrophysiological recordings from tarsal contact chemoreceptors. The glucosinolates present both inside and on the surface of cauliflower leaves were determined. The total amounts obtained with the two methods differed by a factor of 100. The extract of the leaf surface contained about 60 μg per g leaf extracted (gle), the total leaf extract 7.5 mg per gle. The glucosinolate patterns of the two extracts were qualitatively similar, but the ratios of the content of individual glucosinolates showed considerable differences. The D sensilla on segment 3 and 4 of the tarsus of D. radicum females were shown to contain a sensitive receptor cell for glucosinolates. In contrast, the receptor cells of the D sensilla of the other segments did not respond in a dose dependent way to these compounds. The glucosinolate receptors were found to be especially sensitive to glucobrassicin, gluconasturtiin and glucobrassicanapin with thresholds of about 10 −8M to 10 −9M. Large differences (up to two orders of magnitude) were observed among the different glucosinolates. A significant correlation was found between the behavioural discrimination index and the electrophysiological results. But no obvious correlation existed between the chemical nature of the glucosinolate side chain (e.g. indole, aromatic and aliphatic groups), and their stimulatory activity. However, a significant correlation was found between the overall length of the side chain and the biological activity. Although the flies discriminated clearly between model leaves with and without glucosinolates, a clear dose response curve was only obtained for the indole glucosinolate glucobrassicin. Since the most stimulatory fraction of the surface extract contained no glucosinolates, it was concluded that other compounds, in addition to glucosinolates, do play an important role for the stimulation of oviposition.

Tarsal contact chemoreceptors of the black swallowtail butterfly Papilio polyxenes (Lepidoptera: Papilionidae) were stimulated with leaf-surface extracts and ethanolic extracts of whole leaves of a host-plant (Daucus carota) and a non-host (Brassica oleracea). Both leaf extracts evoked large numbers of spikes but stimulated different receptor neurones. It is concluded that a large difference exists between the sensory responses to host and non-host extracts. Two chemicals, luteolin 7-0-(6“-0-malonyl)-β-D-glucopyranoside and trans-chlorogenic acid, known to be present in the host and known oviposi-tion stimulants for P.polyxenes, were also tested and shown to be active. No responses were found to luteolin 7-O-β-D-glucoside or to luteolin 7-O-β-D-glucuronide. These flavonoids occur in D.carota foliage, but do not stimulate oviposition.

According to sympatric speciation theory, adaptation to different host plants is expected to pleiotropically lead to assortative mating, an important factor in the reduction of gene flow between the diverging subpopulations. This scenario predicts mating on and oviposition preference for the respective hosts in both the diverging subpopulations and recently originated species. Here, we test both predictions in the oligophagous Yponomeuta padellus (L.) and the monophagous Yponomeuta cagnagellus (Hubner) (Lepidoptera: Yponomeutidae), two closely related small ermine moth species from the western European clade of Yponomeuta for which speciation in sympatry has been proposed. Mating location and adult host acceptance were evaluated under both semi-field (in a large outdoor cage with a choice of host and non-host plants) and field conditions. In the semi-field experiment, only Y. cagnagellus showed some preference for mating on its own host (16% of all mating pairs) over non-host plants (3% of all mating pairs). However, in both species, more than 80% of the mating pairs were not formed on a plant but instead on the cage itself. Further examination of the mating site of Y. cagnagellus in the field revealed no preference for host plants over non-host plants in the two consecutive years of observation. Yponomeuta padellus females, collected from and reared on Prunus spinosa L. (Rosaceae), showed an oviposition preference for the alternative host Crataegus monogyna Jacq. (Rosaceae) in the semi-field experiment. We thus found no evidence that host-plant fidelity (in terms of mating site) has been the driving force in the speciation process of these Yponomeuta species, nor did we find evidence of host race formation in the tested population of the oligophagous Y. p adellus .

The effects of foliar form, colour and surface characteristics on the oviposition behaviour of the cabbage fly were investigated by presenting females with various leaf models made of paper. The models differed in shape, size, colour, surface coating, the presence of 3 dimensional folds and a stem. It was shown that in the presence of host leaf surface extracts physical factors can strongly influence oviposition behaviour. Females laid the most eggs around the base of bright green or yellow models having a stem, vertical folds and covered with a thin layer of paraffin. The vertical folds in the surface increased the chance of a behavioural transition from leaf run to stem run. Zusammenfassung Eiablageverhalten der Kohlfliege beeinflusst durch Blatt-Form, Farbe und Oberflachestructur Mit Hilfe von Papiermodellen von Kohlblattern wurde der Einfluss von visuellen und mechanischen Eigenschaften des Blattes auf das Eiablageverhalten der kleinen Kohlfliege, Delia radicum, untersucht. Die verwendeten Modelle unterschieden sich in Form, Grosse, Farbe, Oberflachen struktur und Stiel des Blattes. Es wurde gezeigt, dass physikalische Reize zusammen mit den chemischen Stoffe auf der Blattoberflache stimulierend wirken. Ein naturliches Eiablageverhalten und die starkste Eiablage wurde durch ein Modell ausgelost, welches sich durch leuchtend grune Farbe, Stiel und vertikale Falten und einen Paraffinnuberzug auszeichnete.

In lepidopterous larvae the maxillary palps contain a large portion of the sensory equipment of the insect. Yet, knowledge about the sensitivity of these cells is limited. In this paper a morphological, behavioral, and electrophysiological investigation of the maxillary palps of Yponomeuta cagnagellus (Lepidoptera: Yponomeutidae) is presented. In addition to thermoreceptors, CO(2) receptors, and gustatory receptors, evidence is reported for the existence of two groups of receptor cells sensitive to plant volatiles. Cells that are mainly sensitive to (E)-2-hexenal and hexanal or to (Z)-3-hexen-1-ol and 1-hexanol were found. Interestingly, a high sensitivity for benzaldehyde was also found. This compound is not known to be present in Euonymus europaeus, the host plant of the monophagous Yponomeuta cagnagellus, but it is a prominent compound in Rosaceae, the presumed hosts of the ancestors of Y. cagnagellus. To elucidate the evolutionary history of this sensitivity, and its possible role in host shifts, feeding responses of three Yponomeuta species to benzaldehyde were investigated. The results confirm the hypothesis that the sensitivity to benzaldehyde evolved during the ancestral shift from Celastraceae to Rosaceae and can be considered an evolutionary relict, retained in the recently backshifted Celastraceae-specialist Y. cagnagellus.

Changes in host acceptance is an important factor in the host specialization of phytophagous insects, and knowledge of the genetic organization of this behaviour is necessary in order to understand how host shifts occur. Here we describe the inheritance of adult host acceptance (oviposition) in three closely related species of Yponomeuta Latreille (Lepidoptera: Yponomeutidae), and their interspecific hybrids. Yponomeuta cagnagellus (Hubner), a specialist on Euonymus europaeus L. (Celastraceae), Y. malinellus Zeller, a specialist on Malus spp. (Rosaceae), and Y. padellus (L.), oligophagous on a number of Rosaceae, were tested for their acceptance of parental hosts in choice tests. Acceptance of E. europaeus is semi-dominant in hybrids of Y. cagnagellus x Y. padellus, and in hybrids of Y. cagnagellus x Y. malinellus. The dominance of this acceptance was confirmed in oviposition tests with backcross hybrids: backcross hybrids Fl x Y. cagnagellus oviposited mainly on E. europaeus and Fl x Y. padellus still deposited more than half of their egg masses on E. europaeus. Reciprocal hybrids did not differ in their host acceptance, indicating that the trait is autosomal. We further studied the effect of larval food on adult host acceptance ('Hopkins host selection principle') in split full-sib F1 families. Larval diet influenced oviposition only in one of two hybrid crosses. The F1 hybrid of Y. padellus x Y. cagnagellus, reared on Prunus spinosa L., deposited a significantly lower percentage of egg masses on E. europaeus compared to their full-sib sisters fed with E. europaeus. We did not find this in the reciprocal cross. However, still more than half of the egg masses are deposited on E. europaeus by hybrids that have no experience on this host. We conclude that the semi-dominant character of acceptance of E. europaeus and a tendency of Rosaceae-feeding Yponomeuta to deposit egg masses on this host may have created the opportunity for the host shift of the predecessor of Y. cagnagellus from Rosaceae to the Celastraceae. This shift may have been further facilitated by a weak tendency of adults to oviposit on their larval food source.

A novel type of locomotion compensator was designed and tested for its use in orientation behavior experiments with a predatory mite. In this apparatus, displacements of the test animal in the two-dimensional plane are recorded using video equipment and a servosphere that keeps the animal in focus. The x and y displacements are registered using two rotation encoders and are compensated using a pair of servo-motors, in such a way that the animal is always positioned on top of the sphere, yet moves freely. Well-fed and starved predators were tested for their responses to (1) still air, (2) a stimulus-free air flow, (3) an air flow with odors from uninfested Lima bean leaves, and (4) an air flow with odors from Lima bean leaves infested by plant-feeding mites, the prey of the predatory mites. Anemotactic responses of adult Phytoseiulus persimilis females were feeding state dependent. Well-fed predators moved downwind under Treatments 1-3 but moved neither up- nor downwind in the presence of odors from infested plants (Treatment 4). Starved predators moved upwind under all treatments. These results are in agreement with those of earlier studies in a wind tunnel, and therefore, the new type of locomotion compensator (LC-100) offers an excellent method for studying the orientation behavior of micro-arthropods.

The cellular localisation of the sensitivity to two host-plant specific larval phagostimulants was determined in Yponomeuta evonymellus and in inter-specific hybrids of Y. cagnagellus and Y. padellus. The combined results of cross-adaptation experiments, analysis of mixture responses and inter-spike interval distributions indicated that the sensitivity for sorbitol and dulcitol is localised in the same cell in the investigated species. This result suggests that during the evolution of the genus the main alteration in the mechanism of sugar-alcohol detection occurred at the level of the receptor proteins. Relatively simple modification of this kind can, however, have large effects, since larvae with modified receptor proteins will directly respond to the new stimulus while all central processing and resulting behaviour can remain unmodified.

The behavioural phase state and coloration of hatchling Schistocerca gregaria were examined in a series of experiments to determine the means by which phase characteristics are passed between generations. Both crowding of solitary-reared females at the time of oviposition and high egg pod densities promoted behavioural gregarization, although the former appeared to be a rather more potent factor. In contrast, egg pod density alone appeared to promote the development of hatchlings with dark patterns characteristic of the gregarious phase. The phase characteristics of hatchlings were unaffected when sand previously used for oviposition was used to collect further egg pods. Early separation of presumptive gregarious eggs from egg pods laid by crowd-reared females led to solitarization of the hatchlings, indicating that a factor, either in or around the eggs, removed by early separation promoted gregarization. Both the eggs and foam plugs of egg pods from crowd-reared, gregarious females appeared to be a source of this gregarizing factor. In contrast, there was no evidence for a solitarization factor in egg pods from solitary-reared S. gregaria. Saline extracts of egg pod foam plugs produced an active factor which promoted gregarization both in eggs from solitary-reared females and in eggs from gregarious females which were separated and washed to removed the factor. Solitary eggs were influenced by the gregarizing factor in foam plug extracts for up to 1 day after oviposition. Saline extracts of foam plug retained their activity for up to 1 day. Initial studies on the properties of this factor were made. We conclude that the foam plugs of egg pods from crowd-reared, gregarious locusts contain a small (

The nature of stimuli, emanating from other locusts, which are effective in inducing gregarization in the desert locust was investigated. Isolated-reared fifth-instar nymphs were subjected to tactile, visual and olfactory stimuli, presented singly and in combination, and the effect on the behavioural phase state was quantified using logistic regression analysis. Tactile stimulation provided by rolling paper spheres proved to be highly gregarizing, whether presented alone or in combination with the other stimuli. Olfactory and visual stimuli together caused partial behavioural gregarization. Visual stimulation alone was weakly gregarizing after prolonged exposure, while olfactory stimuli alone were ineffective. Nymphs and pre-reproductive and reproductive adults of both sexes were also treated with synthetic adult male `aggregation' pheromone blend (Torto et al., 1994, Journal of Chemical Ecology 20, 1749). No effect of this blend was found on the behavioural phase state, even when visual stimuli were present. Non-locust related stimuli, including wheat odour and flashing lights, were also tested on nymphs. Neither induced any change in the behavioural phase state, indicating that increased sensory flow is not a sufficient explanation for locust-induced behavioural phase change.

Oviposition by the cabbage root fly, Delia radicum (L.) (Dipt., Anthomyiidae) is governed to a large extent by chemicals present on the leaf surface of the host plant Brassica oleracea L. Although this fact has been recognized for some time, knowledge about the nature of the phytochemicals is still far from complete. Glucosinolates (mustard oil glucosides) and their breakdown products, the isothiocyanates (mustard oils), have been known to be involved. The glucosinolates stimulate oviposition and the tarsal D-sensilla contain chemoreceptors sensitive to this group of compounds (Stadler, 1978; Roessingh et al., unpubl.).

The role and nature of volumetric feedbacks in the regulation of meal size was investigated for Schistocerca gregaria. Feedback from the anterior region of the crop was found to be important and denervation led to hyperphagia. Additional feedbacks were found from more posterior gut regions. Cannulating agar or paraffin into the mid- and hindgut caused a decrease in the size of a following meal. This effect was removed by cutting the ventral nerve cord anterior to the terminal abdominal ganglion. Increasing the blood volume by injections of isotonic saline into the haemocoel was also found to decrease subsequent meal size. This effect was not mediated via the gut or body wall stretch receptors and possible mechanisms are discussed. The results for S. gregaria are compared with those obtained previously from Locusta migratoria. RESUME Feed back volumetrique et controle de la taille du repas chez Schistocerca gregaria L'etude de la nature et du role des feed backs volumetriques dans la regulation de la taille du repas a ete effectuee chez S. gregaria. Un feed back, originaire de la region anterieure du jabot, s'est revele important et la denervation a provoque l'hyperphagie. Des feed backs supplementaires ont ete deceles dans des regions plus posterieures du jabot. De l'agar ou de la paraffine canules dans la jabot moyen et posterieur ont provoque une diminution de la taille du repas suivant. Ces effets sont supprimes avec la section de la corde nerveuse centrale avant le ganglion abdominal terminal. L'augmentation du volume sanguin par des injections de solution saline isotonique a aussi diminue la taille du repas suivant. Cet effet n'est pas transmis via des recepteurs de tension du jabot ou de la paroi du corps; des mecanismes envisageables sont discutes. Ces resultats avec S. gregaria sont compares a ceux obtenus anterieurement avec Locusta migratoria.

In this study the trail following behaviour of the caterpillar Yponomeuta cagnagellus (Hubner) (Lepidoptera, Yponomeutidae) is investigated. It is demonstrated that these caterpillars follow trails made by conspecifics. Ablation experiments show that at least part of this behaviour is directed by the tactile senses, but additional chemical cues cannot be excluded. In choice experiments using trails from different species, Y. cagnagellus strongly preferred conspecific trails over those from Malacosoma neustria, but did not prefer conspecific over other Yponomeuta trails. This lack of species-specificity within Yponomeuta is discussed and it is concluded that trail following is probably of little help in the elucidation of the evolutionary history of the genus. Resume La facon dont Ypotiomeuta cagnagellus suit les pistes Cette etude traite de la facon dont les chenilles de Y. cagnagellus Hubner (Lepido. Yponomeutidae) suivent des pistes. Elles suivent les pistes de leurs congeneres. Par ablation, on peut montrer qu'une partie au moins de ce comportement est due a des perceptions tactiles, mais que l'intervention d'informations chimiques complementaires ne peut etre exclue. En presence de choix entre traces de differentes especes, elles preferent nettement les traces des congeneres a celles de Malacosoma neustria, mais ne choisissent pas entre les differentes especes d'yponomeutes. L'absence de specificite parmi le genre Yponomeuta montre que la selection des pistes est probablement de peu d'utilite pour l'etude de l'histoire evolutive du genre.

. Caterpillars of the genus Malacosoma follow trails of the chemical 5-beta-cholestane-3,24-dione, but nothing is known of how they perceive this compound, or more generally about the sensory basis of trail following in caterpillars. By selective ablations of chemosensory organs we show that, in Malacosoma, the trail chemical is perceived by the maxillary palpi. In another lepidopteran species, Yponomeuta cagnagellus, the palpi are needed to discriminate their own trails from a trail of Malacosoma. Malacosoma larvae also lose their specificity for conspecific trails when their palpi are ablated. Volatile cues evidently do not play a role in trail-following behaviour, since neither Malacosoma nor Yponomeuta can orient on a trail covered with fine nylon mesh. These data indicate that for Malacosoma, and probably also for Yponomeuta, contact chemoreception mediated by the maxillary palpi is the primary mode of pheromone perception. The evolution of receptor sensitivity to trail chemicals in caterpillars is discussed.