A Survey of Husbandry Practices for Lorisid Primates in North American Zoos and Related Facilities

Zoos and related facilities in North America currently manage five species in the primate family Lorisidae: the greater (Nycticebus coucang), Bengal (N. bengalensis) and pygmy (N. pygmaeus) slow lorises, red slender loris (Loris tardigradus), and potto (Perodicticus potto). We used an online survey to describe institutional housing and husbandry practices for these species and assess the extent to which practices are consistent with established guidelines. Our results show that most captive lorisids are housed solitarily or in pairs. Most individuals occupy a single exhibit space in a building dedicated to nocturnal animals. Facilities are commonly meeting recommendations for abiotic exhibit design and are providing animals with an enriched environment. However, pottos and slender lorises currently occupy exhibit spaces smaller than the recommended minimum, and the impact of cleaning protocols on olfactory communication should be critically evaluated. Few facilities are taking advantage of the benefits of positive reinforcement training for promoting animal welfare. Research is greatly needed on the effects of exhibit lighting on behavior, health, and reproduction; and to determine how best to manage the social needs of lorisids with naturally dispersed social structures. Although captive populations of slender lorises, pottos, and slow lorises are declining, we suggest that improved husbandry knowledge has the potential to positively influence population sustainability and to enhance future efforts to manage the growing pygmy loris population. Zoo Biol. 32:88‐100, 2013. © 2012 Wiley Periodicals, Inc.     

Responses of adult Hydrellia lagarosiphon to a revised diet: implications for life cycle studies and laboratory culturing techniques

Preservation of an insect culture under laboratory conditions is essential for its study. Numerous diets have been developed for entomophagous insects undergoing screening as biological control agents in attempts to improve the nutritional quality of food provided in laboratory settings. However, less emphasis has been given to developing a more effective laboratory diet for phytophagous insects with the adult life stage not dependant on the target species. The larvae of Hydrellia lagarosiphon Deeming (Diptera: Ephydridae) mine the leaves of Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae). This species is currently under consideration as a biological control agent of this aquatic invasive. Rearing techniques for the adult stage of other Hydrellia species have been developed but current diets are not ideal as they result in relatively low reproduction rates. We compared alternative nutritional regimes and quantified their impact on life history attributes of H. lagarosiphon. The diets included the previously developed yeast‐sugar diet, a newly developed insect‐derived diet, and a diet that combined the two. Total fecundity was significantly higher for females on an insect‐derived diet compared with the traditional carbohydrate diet and the net reproductive rate (R_o) was also higher. Population doubling time (T_d) was lower, decreasing by 30% compared to the traditional laboratory diet developed for Hydrellia species. Adult females fed the combination diet, including both insect and non‐insect foods, laid 30% fewer eggs than those reared on an insect diet alone. Consequently, insect‐derived nutritional regimes could improve culturing techniques significantly and if permission to release the agent is granted, this diet may benefit mass rearing efforts potentially saving time and reducing associated costs.     

Diet composition,feed preferences and mouth morphology of early stage silver therapon (Leiopotherapon plumbeus,Kner 1864) larvae reared in outdoor tanks

This study examined the diet composition, feeding preferences, and mouth morphology of the silver therapon (Leiopotherapon plumbeus, Kner 1864) larvae under captive conditions. Larvae were reared in outdoor tanks (4 m³) with natural food grown 2 weeks prior to start of larval rearing. Food preference was measured by the Chesson's electivity index (α_i). Gut content analysis of larvae sampled between 5 and 25 days after hatching (DAH) showed the dominance in the diet by zooplankton, mainly copepod nauplii, cladocerans and insect larvae. Small fish larvae (5–9 DAH; 3.32–6.29 mm standard length) preferred cladocerans, ciliates and copepod nauplii; whereas older larvae (12–25 DAH; 5.45–19.26 mm standard length) preferred insect larvae over cladocerans and adult insects. The mouth gape size at 5 DAH was 359 μm and increased to 3.75 mm at 40 DAH when body size grew at an average rate of 0.59 mm d^?1. The standard length (SL) of L. plumbeus larvae was strongly associated with mouth size (r² = 0.98, P < 0.05), indicating a progressive increase of ingested prey size of the fish larvae. These results clarified the early life feeding ecology of this species, which is essential in developing effective hatchery techniques.     

Slow Lorises (<Emphasis Type="Italic">Nycticebus</Emphasis> spp.) Really Are Slow: a Study of Food Passage Rates

The characteristics of food ingested by a primate affect its assimilation of energy by modulating food passage rate. In general, digestive time increases in folivorous primates and decreases in frugivorous primates when they are fed higher fiber diets but this relationship is understudied in exudativorous primates. We compared the food passage rate of five slow loris species. We studied 34 wild-caught slow lorises (15 Nycticebus coucang, 15 N. javanicus, and 4 N. menegensis) in an Indonesian rescue center and four captive-born slow lorises (2 N. bengalensis and 2 N. pygmaeus) in a UK institution. We fed the Indonesian subjects two different diets: a captive-type diet comprising fruits, vegetables, and insects and a wild-type diet formulated to be similar in nutrients to that consumed by slow lorises in the wild, consisting of gum, insects, vegetables, and nectar. We fed the UK subjects a diet of gum, vegetables, insects, and hard-boiled eggs. We formulated this diet to mimic the wild diet, with notably higher fiber fractions and lower soluble sugars than the previous diet. We measured two variables: the transit time (TT) and the mean retention time (MRT). We mixed 1 tsp. of glitter in bananas or gum as our markers and fed them to the slow lorises immediately before their main diet. We noted the date and time of feeding and of appearances of the marker in feces. We weighed food given and left over for each individual to calculate ingested foods and nutrients. We found that TTs were not affected by diet treatment but MRTs were significantly longer for all species fed the wild-type diet. Our results show that Nycticebus spp. have long MRTs for their body weight, and N. pygmdaeus may have the slowest MRT of all primates in relation to body mass. The digestive flexibility of exudativorous primates should allow them to maximize fermentation opportunities when they ingest more (appropriate) fiber by increasing the amount of time the fiber substrate stays in the large intestine. Exudativorous primates appear to have plastic digestive strategies that may be an adaptation to cope with relatively nutrient-poor staple food sources such as gum. The provision of gum in a captive setting may therefore provide benefits for gut health in slow lorises.     

Plant genotype affects the quality of oilseed rape (Brassica napus) for adults and larvae of the pollen beetle (Meligethes aeneus)

Plant quality is one of the main factors influencing the fitness of phytophagous insects. Plant quality can vary not only among genotypes of the same host plant species, but also relative to the insect sex or its life stage. In the present study, the performance of larvae and adults of the pollen beetle (Meligethes aeneus F., Coleoptera: Nitidulidae), a major insect pest of oilseed rape crops, is compared on six genotypes of oilseed rape (Brassica napus). All of the traits that are measured vary among genotypes, and comprise larval developmental duration, life span of unfed emerging adults and survival time of field‐sampled adults fed with pollen from the different genotypes. No correlation is found between insect performance and quantity of food available, showing that the quality of the food (i.e. pollen) is the fitness determinant for this insect species. Additionally, the performance of larvae and adults is also not correlated despite use of the same plant genotypes, suggesting that the determinants of pollen quality differ at least partially between both life stages. It is hypothesized that this may be a result of extensive differences in diet breadth between the life stages: larvae are specialists of brassicaceous plants, whereas adults are generalists. Finally, it is suggested that the manipulation of plant quality to increase pollen beetle development time may comprise a valuable strategy for favouring biological control by natural enemies of this pest; for example, as a result of extending the vulnerability window of larvae to attack by parasitoids.     

Does host‐feeding on GNA‐intoxicated aphids by Aphelinus abdominalis affect their longevity and/or fecundity?

Transgenic potatoes have been transformed with a gene coding the snowdrop lectin Galanthus nivalis agglutinin (GNA) and they have been shown to be partially resistant to aphids. GNA binds to insect gut cells, including those of aphids, consequently inducing disruption of nutrient assimilation. Aphid parasitoids are important natural biocontrol agents of aphids and some species such as Aphelinus abdominalis are commercially available. Aphid parasitoids are endoparasitoids during their larval stages and free-living insects as adults. They could be directly or indirectly affected by GNA during both these stages. In this work, we present data on the potential direct and indirect effects of GNA on adult A. abdominalis. Aphelinus abdominalis is a synovigenic species (eggs are matured throughout the adult life) which needs a diet relatively rich in proteins and amino-acids to produce anhydropic eggs (large, yolk-rich eggs that do not expand in the host during embryonic stages). Adult A. abdominalis females feed on aphid haemolymph and they may be directly exposed to the entomotoxin or indirectly affected by a change in aphid haemolymph quality due to intoxication by GNA. We conducted a first tier experiment to investigate this potential risk. A. abdominalis females were offered either aphids reared on control diet or aphids reared on GNA 0.1% diet as hosts (i.e., as food and oviposition sites). No trace of GNA was found in females fed with GNA-aphids but no GNA could be detected in the haemolymph of aphids fed a 0.1% GNA diet. Longevity and fecundity were not affected suggesting that the quality of the haemolymph necessary for A. abdominalis egg maturation and production was not significantly altered.     

Editorial note

Generalist herbivores can face many challenges when choosing their host plant. This can be particularly difficult if their choice and performance are affected by host experience. Greenhouse whitefly, Trialeurodes vaporariorum Westwood (Hemiptera: Aleyrodidae), is an invasive generalist herbivore, which has established in year‐round greenhouses at northern latitudes where it cannot overwinter outdoors. It mainly uses crops such as cucumber (Cucumis sativus L.), tomato (Solanum lycopersicum L.), and ornamentals as host plants. However, every summer the insect escapes greenhouses and is exposed to natural vegetation. We evaluated the performance of T. vaporariorum on diverse vegetation outside greenhouses after prolonged experience of greenhouse crops. First, we surveyed the vegetation near infested greenhouses. Development success of the insect differed among wild hosts. We identified five new hosts among 12 plant species that bore pupae and were thus considered suitable as the insect's host plants. Members of the Urticaceae and Onagraceae were the most preferred and frequently inhabited by all insect life stages. The highest abundance of insects occurred in plots with low plant species richness, independent of plant family in these habitats. We then studied experimentally the impact of 1 year of preconditioning to one of three common greenhouse crops, cucumber, tomato, or poinsettia (Euphorbia pulcherrima Willd. ex Klotzsch), on the performance of the preconditioned adults and their progeny on four wild plants. Adults from tomato and poinsettia preferred the novel host species over the species to which they were preconditioned. The whitefly population preconditioned to cucumber was the most fecund on all offered hosts. We conclude that generalist herbivores can have large variation in performance, despite polyphagy, on novel hosts as shown by the variable abundance of T. vaporariorum pupae among outdoor hosts. Furthermore, performance of whiteflies on natural vegetation was affected by experience on greenhouse crops. Based on our observations, we provide insights and recommendations for pest management.     

Offspring diet supersedes the transgenerational effects of parental diet in a specialist herbivore Neolema abbreviata under manipulated foliar nitrogen variability

Diet quality influences organismal fitness within and across generations.For herbivorous insects,the transgenerational effecets of diet remain relatively underexplored.Usinga3×3×2 factorial experiment,we evaluated how N enrichment in parental diets of Neolemd abbreviata(Larcordaire)(C oleoptera:Chrysomelidae),a biological control agent for Tradescantia fluminensis Vell.(Commelinaceae),may influence life history and performance of Fi and F2 offspring under reciprocal experiments.We found limited transgenerational effects of foliar nitrogen variability among life-history traits in both larvae and adults.Larval weight gain and mortality were responsive to parental diet contrary to feeding damage,pupal weight and duration taken to pupate.There were significant parental diet x test interactions in larval feeding damage,weight gain,pupal weight and time to pupation.Generally,offspring from parents under high N plants performed better even under low N test plants.Adult traits including oviposition selection,feeding weight and longevity did not respond to the efects of parental diet nor its interaction with test diet as was the case in the larval stage.However,the main efects of test diet were more important in determining adult performance in both generations suggesting limited sensitivity to parental diet in the adult stage.Our results show conflicting responses to parental diet between larvae and adults ofthe same generation among an insec species with both actively feeding larual and adult life stagee These tranegeneratinonal efferte,or lack thereof,may have implications on the field performance of N.abbrevita under heterogencous nutritional landscapes.     

Divergent immune priming responses across flour beetle life stages and populations

Growing evidence shows that low doses of pathogens may prime the immune response in many insects, conferring subsequent protection against infection in the same developmental stage (within‐life stage priming), across life stages (ontogenic priming), or to offspring (transgenerational priming). Recent work also suggests that immune priming is a costly response. Thus, depending on host and pathogen ecology and evolutionary history, tradeoffs with other fitness components may constrain the evolution of priming. However, the relative impacts of priming at different life stages and across natural populations remain unknown. We quantified immune priming responses of 10 natural populations of the red flour beetle Tribolium castaneum, primed and infected with the natural insect pathogen Bacillus thuringiensis. We found that priming responses were highly variable both across life stages and populations, ranging from no detectable response to a 13‐fold survival benefit. Comparing across stages, we found that ontogenic immune priming at the larval stage conferred maximum protection against infection. Finally, we found that various forms of priming showed sex‐specific associations that may represent tradeoffs or shared mechanisms. These results indicate the importance of sex‐, life stage‐, and population‐specific selective pressures that can cause substantial divergence in priming responses even within a species. Our work highlights the necessity of further work to understand the mechanistic basis of this variability.     

Interactions between phytophagous insects and their Opuntia hosts

Abstract.

• 1 The cactophagous insect community on opuntias is analysed to show the number of insect species in different taxa. An extension of this analysis gives the average species complement on large and small opuntias.
• 2 A highly significant positive correlation is found between the total number of phytophagous insect species on individual Opuntia species and a measure of the overall ‘architecture’ of their host plants.
• 3 The specificity of the phytophagous insects on opuntias is briefly considered and the community as a whole analysed by guilds. The co-evolution of the Opuntia-feeding insects and their hosts has culminated in a community of specialist insects to the exclusion of nearly all generalist phytophages.
• 4 The life history strategies of the Opuntia-feeding insects are reviewed. Common to all developmental stages are morphological and behavioural adaptations that reduce the risk of attack by natural enemies. This is clearly the consequence of living on structurally simple host plants where there is little place to hide.
• 5 The possible influence of insect herbivores on Opuntia evolution is discussed.
• 6 An understanding of the interactions between the phytophagous insect community and opuntias has clear implications for the biological control of alien Opuntia weeds.

     

Skeletal development in Pan paniscus with comparisons to Pan troglodytes

Fusion of skeletal elements provides markers for timing of growth and is one component of a chimpanzee's physical development. Epiphyseal closure defines bone growth and signals a mature skeleton. Most of what we know about timing of development in chimpanzees derives from dental studies on Pan troglodytes. Much less is known about the sister species, Pan paniscus, with few in captivity and a wild range restricted to central Africa. Here, we report on the timing of skeletal fusion for female captive P. paniscus (n = 5) whose known ages range from 0.83 to age 11.68 years. Observations on the skeletons were made after the individuals were dissected and bones cleaned. Comparisons with 10 female captive P. troglodytes confirm a generally uniform pattern in the sequence of skeletal fusion in the two captive species. We also compared the P. paniscus to a sample of three unknown‐aged female wild P. paniscus, and 10 female wild P. troglodytes of known age from the Taï National Park, Côte d'Ivoire. The sequence of teeth emergence to bone fusion is generally consistent between the two species, with slight variations in late juvenile and subadult stages. The direct‐age comparisons show that skeletal growth in captive P. paniscus is accelerated compared with both captive and wild P. troglodytes populations. The skeletal data combined with dental stages have implications for estimating the life stage of immature skeletal materials of wild P. paniscus and for more broadly comparing the skeletal growth rates among captive and wild chimpanzees (Pan), Homo sapiens, and fossil hominins. Am J Phys Anthropol 2012. © 2012 Wiley Periodicals, Inc.     

Selective Feeding of <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> (Hübner) and <Emphasis Type="Italic">Spodoptera litura</Emphasis> (Fabricius) on Meridic Diet with <Emphasis Type="Italic">Bacillus thuringiensis</Emphasis> Toxins

Laboratory experiments were conducted to evaluate the behavior of Helicoverpa armigera (Hübner) and Spodoptera litura (Fabricius) larvae on meridic diet with different concentrations of Bt spray formulation Delfin or isolated Cry1Ac protein or the foliage and bolls from transgenic cotton, Bollgard hybrid RCH-317 Bt. Both insect species selectively fed on nontreated diet compared with the diet treated with Delfin. While H. armigera exhibited concentration response with Cry1Ac, this protein did not affect S. litura larvae. In general Helicoverpa selected diet with low concentrations (EC₂₀ and EC₅₀ levels) of Cry1Ac compared with higher concentrations of Cry1Ac. In order to develop appropriate management strategies, a thorough understanding of the behavioral mechanisms leading to the responses of insects to the proteins in transgenic varieties is required. Thus, based on results of the insects fed individually on the leaf discs or bolls from transgenic cotton plants alone or under choice situation with meridic diet revealed that H. armigera larvae preferred meridic diet to transgenic leaves or bolls expressing Cry1Ac protein. H. armigera larvae preferred meridic diet to plant material; more than 70% larvae were seen on the meridic diet, and average larval weight gain was in the range of 121.7–130.5 mg. However, in case of S. litura the larvae showed no significant discrimination between meridic diet and the leaf discs. In fact more than 60% larvae preferred leaf discs for feeding, though Cry1Ac expression in leaf discs was in the range of 0.9–2.18 μg/g. Thus differences in behavioral response could potentially impact the level of efficacy of crop cultivars that have been genetically engineered to produce these proteins.     

Thermal stress exposure of pupal oriental fruit fly has strong and trait-specific consequences in adult flies

Global climate change is projected to increase the incidence of heat waves, their magnitude and duration resulting in insects experiencing increasing environmental stress in both natural and managed ecosystems. While studies on insect thermal tolerance are rapidly increasing, variation across developmental or juvenile stress cross-stage effects within and across generations remain largely unexplored. Yet in holometabolous insects, heat stress at an early developmental stage may influence performance and survival during later stages. Here, we investigated the effects of pupal mild heat stress on the performance of laboratory-reared adult Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) measured as longevity, critical thermal maximum (CT_max), critical thermal minima (CT_min), heat knockdown time (HKDT) and chill coma recovery time (CCRT). Pupal heat stress significantly influenced performance of B. dorsalis adults resulting in impaired longevity and heat tolerance (CT_max and HKDT) in both sexes with improved and compromised cold tolerance (CT_min and CCRT) in females and males, respectively. These findings highlight the role of juvenile stages in mediating stress responses at adult stages. For B. dorsalis, pupal heat stress largely compromised thermal tolerance implying that the species has limited potential to shift its geographic range in heat prone areas. Significant benefits in cold tolerance in females following heat stress may help in improving survival in the cold in the short-term despite restricted activity to the same traits in males. This study suggests that basal heat tolerance and not short-term compensatory thermal plasticity following heat stress may have aided the recent invasion of B. dorsalis in African landscapes.     

The Role of Fruits and Insects in the Nutrition of Frugivorous Bats: Evaluating the Use of Stable Isotope Models1

We estimated the relative contribution of fruits and insects as sources of dietary protein in two species of Neotropical frugivorous bats (Artibeus jamaicensis and Sturnira lilium) using stable carbon and nitrogen isotope analyses. An insectivorous species (Pteronotus parnellii) was also included for comparison. We found constant patterns in stable carbon and nitrogen isotope composition in blood that separated the two species of frugivorous bats from the insectivorous bat. When we used these isotopic values (combined with those of dietary fruits and insects) to estimate the percent contribution of fruits and insects to the diet of the bats, we obtained different results, depending on assumptions and model adopted. We tested models using both 8“N and 8′³C results simultaneously and separately and further used diet‐tissue fractionation factors of 3%o for nitrogen and 1 and 3.5%o for carbon. We found that a carbon‐based model with a diet‐blood enrichment factor of 3.5%o produced the most parsimonious results. The model estimated that A. jamaicensis and S. lilium obtained most of their protein requirements from fruits, whereas P. parnellii fed mostly on insects. No sexual or seasonal variations in the diet of the two frugivorous species were detected. We found no evidence that the diet of sexually active females differed from that of nonsexually active females in the two species of frugivorous bats. We suggest that future studies better define isotopic fractionation between diet and tissues of bats using captive rearing and controlled diets.     

Developmental O-glycan profile analysis shows pentasaccharide mucin-type O-glycans are linked with pupation of Tribolium castaneum

Eukaryotic cells can decorate their proteins with carbohydrate structures or glycans, significantly affecting the properties and activities of these proteins. Despite the importance of protein glycosylation in numerous biological processes, our knowledge of this modification in insects is far from complete. While N-glycosylation is the most studied, the study of O-glycans in insects is still very fragmentary and these studies are limited to a specific developmental stage or a specific tissue. In this article, matrix-assisted laser desorption/ionization (MALDI)-Fourier transform ion cyclotron resonance (FTICR) mass spectrometry (MS) technology was used to analyze the O-glycan profile for the different developmental stages of egg, larva, pupa, and adult of the red flour beetle Tribolium castaneum, an important insect model and pest worldwide. The results on the O-glycan profile showed that the mucin-type glycans dominate the O-glycome of the red flour beetle. Interestingly, some of the more complex mucin-type O-glycans, such as a tetra- (O-GalNAcGalGlcAGalNAc) and pentasaccharide O-glycan (O-GalNAc(GalGlcA)GalNAcGlcA), were highly abundant during the pupa stage, the intermediate stage between larval and adult stage in holometabolous insects, demonstrating that insect metamorphosis is accompanied with a change in the insect O-glycan profile. Together with the N-glycan profile, the current data are a foundation to better understand the role of protein glycosylation in the development of insects.     

Ontogeny of protein concentration,haemocyte concentration and antimicrobial activity against Escherichia coli in haemolymph of the invasive harlequin ladybird Harmonia axyridis (Coleoptera: Coccinellidae)

The harlequin ladybird is considered to be one of the most successful invasive insect species. Among other traits, its invasive success is considered to be caused by a powerful immune system. In the present study, we investigate the ontogenetic profile of protein concentration, concentration of circulating haemocytes and constitutive antimicrobial activity against Escherichia coli in Harmonia axyridis haemolymph during late larval development and early adult life. Protein concentration increases during the first 32 days of adult life from 45 to 100 mg per mL of haemolymph and reaches intermediate values during larval stages. The concentration of circulating haemocytes is very low (5000 haemocytes per μL of haemolymph) in late larval stages and increases strongly during first 8 days of adult life to values of approximately 30 000 haemocytes per μL of haemolymph. The killing efficiency of haemolymph against E. coli is lowest in larval stages, rapidly increases in the prepupal stage and then steadily grows during the whole period of adult life. There are no significant effects of sex on any of the investigated physiological or immune parameters. In general, the patterns observed for H. axyridis contrast with many results that are reported for other insects (e.g. bees, fruit flies, crickets or mosquitoes). One possible explanation is the contrasting life history of H. axyridis, with a fast preimaginal development and a long adult lifespan being linked to a long reproductive period. Substantial variation in physiological and immune parameters during ontogeny also has important methodological implications because individuals of exactly the same stage/age have to be employed for comparative studies.     

Diet and Feeding Behavior of Mysore Slender Lorises

We studied the feeding ecology of the Mysore slender loris (Loris lydekkerianus lydekkerianus) for 10.5 mo in a dry scrub forest at Ayyalur Interface Forestry Division, Tamil Nadu, South India. We recorded and analyzed 1240 feeding incidents, which indicate that the lorises were almost exclusively faunivorous, with 96% of all feeding events representing animal prey. Of prey items that could be identified (n = 605), 62.9% were ants and termites. Lorises fed on 9 orders and 17 families of insects, plus spiders, molluscs, and small vertebrates. Lorises infrequently fed on gums and a legume pod. They usually grabbed prey with one hand, while other appendages firmly held the substrate. Many of the identifiable prey items belong to insect taxa likely to contain toxic chemicals. Consumption of insects inferred to be toxic was accompanied by an elaborate behavioral repertoire of sneezing, slobbering and urine-washing. A high proportion of insects eaten by slender lorises (71%) occurred in patches or aggregations. The utilization of aggregated social insects may have implications for understanding the unusually high degree of gregarious behavior exhibited by the lorises.     

Comparative feeding selectivity of herbivorous insects on water lilies: aquatic vs. semi‐terrestrial insects and submersed vs. floating leaves

1. The rate of grazing damage experienced by submersed and floating leaves of water lilies (Nuphar variegata and Nymphaea odorata) was monitored in lakes in the Upper Peninsula of Michigan, U.S.A. Herbivores damaged 0.2–1.7% of the leaf surface of water lilies per day. These grazing rates differed between plant species, between submersed and floating leaves, and between lakes. Some leaves had more than 60% of their surface damaged and an overall mean of 16% damage occurred during the 2–3 week monitoring period of this study. 2. Snapshot measurements of grazing damage on randomly collected submersed and floating leaves of Nuphar showed that submersed leaves were more damaged (11.0 ± 1.6%, n = 84) than floating leaves (3.8 ± 0.6%, n = 92). Overall, these 176 Nuphar leaves had 7.2% of their area damaged. 3. Five species of herbivorous insects were commonly found on water lilies (Nymphaeacea). One primarily aquatic insect (sensu 1 ), a caddisfly larva (Trichoptera: Limniphilidae), had a generalized diet of water lilies, other macrophytes, algae, and detritus. Four of the five insects were from primarily terrestrial insect groups (Coleoptera and Diptera;‘secondary invaders’, sensu 1 ) and consumed only water lilies in food preference experiments. 4. The feeding preferences of the generalist trichopteran were altered when the macrophytes were freeze-dried, ground into a powder, and reconstituted in an alginate gel. This suggests that plant structure may be an important feeding determinant for this insect. In contrast, a specialist weevil preferred its host plant in choice assays, regardless of whether fresh tissue or reconstituted macrophytes were used, suggesting this insect cued on a unique, non-structural property of its host plant. 5. These results suggest that herbivory on freshwater macrophytes is of a similar magnitude to that on terrestrial plants. The findings of this study are consistent with the hypothesis that herbivorous insects of primarily terrestrial groups have a narrower diet breadth than insects of primarily aquatic groups.     

Effect of diet on the fecundity of three carabid beetles

Abstract In the present study, the importance of diet in terms of fecundity is compared for three species of the carabid genus Amara (Coleoptera: Carabidae), using an insect diet, two types of seed diet (Capsella bursa‐pastoris, Stellaria media) and a mixed diet. It is expected that the species of carabid studied have different food requirements for reproduction. Diet affects reproduction performance and egg production significantly. A mixed diet and both single‐seed diets are suitable for reproduction in Amara aenea (DeGeer) because a higher proportion of the females reproduce and lay significantly more eggs than on a purely insect diet. Females of Amara familiaris (Duftschmid) do not reproduce unless provided with seeds of S. media. Seeds of C. bursa‐pastoris or a mixed diet are equally suitable diets for reproduction of Amara similata (Gyllenhal); a diet of insects or seeds of S. media is unsuitable. The results support the hypothesis that the species under investigation have specific food requirements, suggesting that seed feeding has evolved to different degrees in particular species: A. aenea is omnivorous, whereas A. familiaris and A. similata specialize on the seed of a particular plant species or family. This resource partitioning facilitates co‐occurrence of carabid species.     

Life history shifts and alterations in the early development of parasitic wasps

Summary

In insects, the regulation of embryonic development has been intensively studied in model species like Drosophila melanogaster. Previous comparative studies have suggested that the developmental processes documented in Drosophila well describe embryogenesis of holometabolous insects generally. However, there have been few attempts to take into account how life history has influenced insect embryogenesis or to characterize early development of species with life histories fundamentally different from flies. Our studies of advanced insects in the order Hymenoptera suggest that punctuated shifts in life history can profoundly influence these events. In particular, alterations associated with the evolution of endoparasitism argue that departures from the fly paradigm may occur commonly among insects that develop under conditions different from typical terrestrial species.