Changes in latitude: overwintering survival of two Lilioceris cheni (Coleoptera: Chrysomelidae) biotypes in Florida

Lilioceris cheni is a successful biological control agent on the invasive yam, Dioscorea bulbifera in many areas in Florida, Georgia, Alabama and Louisiana. Two biotypes are available for release and come from significantly different centres of origin. The Nepalese biotype was collected at higher altitude and latitude sites, whereas the Chinese biotype was collected at slightly lower latitude and lower altitude. We hypothesised that Nepalese beetles would survive winter better in north Florida and that Chinese beetles would overwinter better in subtropical south Florida. We established sites in Homestead, Fort Pierce, Gainesville and Tallahassee, Florida to gauge the overwintering survival of each biotype. Beetles that overwintered in each site for one to three seasons were collected and followed to assess their reproductive capacity. In the final year of the experiment, we extracted fat bodies to determine effects of biotype and site. Nepalese beetles overwintered significantly better overall. Beetles survived better in the first study year (2013–2014). Both biotypes reproduced very well after surviving winter and beetles in Fort Pierce survived better than beetles in other regions and maintained a higher fat body. Nepalese beetles may have a wider ecological envelope – a decided advantage for a large region such as peninsular Florida. We suggest that current release programmes incorporate Nepalese beetles into their protocol to increase overwinter survival and establishment.     

Measuring success of a reintroduced population of the American burying beetle (<Emphasis Type="Italic">Nicrophorus americanus</Emphasis> Olivier) to Nantucket Island,MA

The American burying beetle (Nicrophorus americanus) is a federally listed endangered beetle and since 1993 multiple organizations have collaborated to reintroduce this species to Nantucket Island, Massachusetts, USA. We present evidence that despite very successful reintroduction methods, the reintroduced population is not self-sustaining and requires human assistance for long term maintenance. Beetles were reintroduced from captive stock and each year we augmented the population by trapping wild beetles, pairing males and females, and supplying the pairs with carrion. Long term monitoring of this population has shown that, when provided with carrion, N. americanus on Nantucket have an over winter survival rate of 15?% and a reproductive success rate of 54?%. After seeing the number of beetles captured between 2007 and 2011 double, we modified protocols to determine if the established population would be self-sustaining and we have seen a drastic decline. We suggest that a lack of natural carrion is the main reason for this decline.     

Standardized capture rates of the endangered American burying beetle, Nicrophorus americanus Olivier (Coleoptera: Silphidae) using different trap protocols

The U.S. Fish and Wildlife Service has adopted two different pitfall trap protocols to survey the endangered American burying beetle, Nicrophorus americanus. One protocol uses a transect of eight pitfall traps that prohibit bait contact over the course of three trap nights. The other protocol uses buckets that allow for bait contact over a five night trapping period. A trap night is defined as one trap open for one night, and the transect protocol has historically been calculated as eight trap nights per calendar night while; the bucket protocol has been calculated as one trap night per calendar night. This study examined the effectiveness of each protocol based on the number of beetles (Nicrophorus spp.) captured per trap night (BTN) in field and laboratory trials. When each transect was considered as a single trap instead of eight separate traps, no significant difference in BTN was detected between the protocols in any year. Laboratory trials were conducted using Nicrophorus marginatus to determine differences in capture efficiency based on protocol, time after release, and feeding status. The proportion of beetles captured after 5 days was greater when compared to 3 days for either protocol. Our results indicate that cup transects used to sample the American burying beetle should be considered as a single trap when calculating trap night and that the use of five trap nights rather than three would increase the likelihood of capturing beetles.     

The effect of soil depth and exposure to winter conditions on survival of the potato tuberworm,Phthorimaea operculella

The potato tuberworm, Phthorimaea operculella Zeller (Lepidoptera: Gelechiidae), is an important pest of solanaceous crops including potato [Solanum tuberosum L. (Solanaceae)]. Recent evidence of survival of the pest in the Columbia Basin of Oregon and Washington, USA, is surprising as potato tuberworm is most commonly a pest in tropical and subtropical regions. Potato tuberworm was studied in a manipulative experiment to determine how stage (egg, larva, and pupa) and soil depth affected the potential for winter survival. In early January 2007, eggs, tubers infested with larvae, and pupae were placed in the soil at 6, 10, and 20 cm depths, or left on the soil surface. Each potato tuberworm life stage was sampled at 7–17‐day intervals for 90 days and survival was estimated at each time period. Eggs survived up to 54 days with the greatest survival observed at 6‐cm soil depth and the lowest survival on the soil surface. However, egg survival was significantly reduced after 1 month of exposure to winter conditions. Larvae were able to survive up to 30 days with the greatest survival observed at 20‐cm soil depth. Tubers at the surface and buried at 6 cm were frozen; thus, no larval survival was recorded. The pupal stage showed a greater tolerance to winter conditions than the egg or larval stages, surviving up to 91 days of exposure. These results suggest that the pupal stage is probably the stage most likely to survive winter conditions in the northwestern USA.     

Impact of cold on the immune system of burying beetle,Nicrophorus vespilloides (Coleoptera: Silphidae)

Insect overwintering is one of the most astonishing phases of the insect life cycle. Despite vast amounts of knowledge available about the physiological mechanisms of this phenomenon, the impact of stress factors on insect immune system functioning during the winter is still unknown. The aim of this study is to analyze how low temperatures influence the immune system of the beetle Nicrophorus vespilloides. The results show that the beetle's immune system is differently modulated by cold induced in laboratory settings than that which occurs in natural conditions. Among beetles cultured in conditions similar to summer, low temperatures, did not influence the number of circulating haemocytes, phenoloxidase activity, haemocytes morphology, and percentage ratio of haemocyte types. In these beetles, differences were noted only in the ability of haemocytes to perform phagocytosis. Individuals acclimated in natural conditions in autumn had a higher level of humoral response and a different percentage ratio of haemocyte types. During the winter period, the number of haemocytes in the beetles decreased, but the percentage ratio of phagocytic haemocytes increased. Furthermore, we noted an increase of phenoloxidase activity. Our study also showed mitotic divisions of haemocytes in haemolymph collected from burying beetles after cold exposure and from burying beetles collected from natural conditions during autumn and winter. Differences in response to low temperatures in laboratory conditions and the natural environment suggest that the simultaneous presence of other stress factors during winter such as desiccation and starvation have a significant influence on the activity of burying beetle's immune system.     

Increased Frequency of Intergroup Encounters in Wild Bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>) Around the Yearly Peak in Fruit Abundance at Wamba

Intergroup interactions in primates vary from nonagonistic to severely aggressive. Food resources and fertile females cause intergroup aggression when groups defend resources and mates from other groups. Peaceful intermingling during intergroup encounters is rare but has been reported in several primates, including bonobos (Pan paniscus). Although intergroup encounters in bonobos occur at both nonprovisioned and provisioned sites, provisioning may be one factor responsible for frequent intergroup encounters. We studied intergroup encounters between one bonobo group (PE) and its neighboring groups, one semihabituated and two habituated groups, under nonprovisioned conditions in 2010–2015 at Wamba, Democratic Republic of Congo. We examined whether fruit abundance and females with sexual swellings influenced encounter frequency. PE group encountered the other groups on 7.1 days (mean, range: 0–19) per month. Up to four different groups met simultaneously at one location. Encounter frequency was highest around the yearly peak in fruit abundance, suggesting that reduced food competition is linked to increased encounter frequency. During periods of relatively low fruit abundance, the probability of an encounter was higher when a higher number of adult females with maximum sexual swellings were present, suggesting that the attractiveness of females with sexual swellings promoted group encounters during periods of relatively low fruit abundance. These findings suggest that both competition for food and the availability of females with sexual swellings influence encounter frequency in bonobos.     

Reduced splash dispersal of bovine parasitic nematodes from cow pats by the dung beetle Diastellopalpus quinquedens.

From a thoroughly mixed portion of cattle feces with Cooperia sp. eggs, 1-kg artificial pats were placed in 6 buckets containing 6 kg of soil each. Ten dung beetles, Diastellopalpus quinquedens, were added to each of 3 buckets. The remaining 3 buckets served as controls without beetles. When infective parasite larvae (L3) had developed in the cow pats indoors, the following procedure was followed. During occasions of rainfall each bucket was placed outdoors in the center of a wider and higher container. When the rain stopped all buckets were brought indoors, and infective larvae spread by splash droplets during the rain were collected in the containers and counted. After 33 days, the remaining dung on the soil surface in buckets with dung beetles constituted only 38% of that in the controls. Moreover, the number of L3 in feces left on the soil surface in the buckets with beetles was reduced by 88%, presumably due to beetle activity. This may explain the 70-90% reduction in splash dispersal of L3 of Cooperia sp. from cow pats attacked by beetles. The dung-burying activity of the beetles did not result in increased numbers of L3 in the soil under the cow pats, suggesting that many of the parasites in buried feces were destroyed.     

Pre-ovipositional maternal care alleviates food stress of offspring in the flower beetle <Emphasis Type="Italic">Dicronocephalus wallichii</Emphasis>

Unlike most other flower beetles, females of Dicronocephalus wallichii exhibit nesting behaviour. The female constructs a burrow in the soil, cuts dead plant leaves into small pieces to provision the nest, and then lays one egg inside the nest. Hatched larvae have been thought to feed on the nest materials prepared by their mothers, but the effects of pre-ovipositional care on larval performance have not been tested. The hatched larvae were found to stay in the nest for 15–30 days until they consumed the nest materials. We examined whether the presence of provisioned nests enhanced larval performance under both benign and food-stress conditions. With high-nutrient soil, larval survival rate and growth speed were not affected by the presence of provisioned nests. By contrast, with low-nutrient soil, mortality of the larvae was much higher in the absence than in the presence of provisioned nests. The growth speed of larvae with nests located in low-nutrient soil was as high as those reared in high-nutrient soil. These results indicate that females alleviate the food stress of larvae during their initial developmental stage by constructing provisioned nests.     

Phoretic Poecilochirus mites specialize on their burying beetle hosts

Recurring species interactions can cause species to adapt to each other. Specialization will increase the fitness of symbionts in the coevolved association but may reduce the flexibility of symbiont choice as it will often decrease fitness in interactions with other than the main symbiont species. We analyzed the fitness interactions between a complex of two cryptic mite species and their sympatric burying beetle hosts in a European population. Poecilochirus mites (Mesostigmata, Parasitidae) are phoretic on burying beetles and reproduce alongside beetles, while these care for their offspring at vertebrate carcasses. While Poecilochirus carabi is typically found on Nicrophorus vespilloides beetles, P. necrophori is associated with N. vespillo. It has long been known that the mites discriminate between the two beetle species, but the fitness consequences of this choice remained unknown. We experimentally associated both mite species with both beetle species and found that mite fitness suffered when mites reproduced alongside a nonpreferred host. In turn, there is evidence that one of the beetle species is better able to cope with the mite species they are typically associated with. The overall fitness effect of mites on beetles was negative in our laboratory experiments. The Poecilochirus mites studied here are thus specialized competitors or parasites of burying beetles.     

Distribution and survival of overwintering adults of the Dutch elm disease vector, Hylurgopinus rufipes (Coleoptera: Scolytidae), in American elm trees in Manitoba

Abstract 1 The native elm bark beetle, Hylurgopinus rufipes, is the principal vector of Dutch elm disease in Manitoba, Saskatchewan and North Dakota, and disease management measures include applying residual insecticides to the lower part of elm tree trunks where the adult beetles overwinter. 2 In American elm trees in southern Manitoba, we counted entrance holes produced by beetles then felled and dissected trees to determine numbers of tunnels and numbers and survival of overwintering beetles. 3 Densities of entrance holes, tunnels and beetles followed a logistic relationship with tree trunk diameter; densities were near zero at diameters < 10 cm and reached a site‐specific asymptote at diameters > 20 cm. 4 Asymptotic densities of holes, tunnels and beetles in samples from 55 to 190 cm above the ground were, respectively, 22%, 22% and 0.7% of those within 25 cm of the ground. 5 Within the height range 0–190 cm, the proportion of living beetles declined steeply with increasing height. 6 Average density of holes at height 0–25 cm estimated from a sample of several trees of diameter ≥15 cm could be used to predict the asymptotic maximum density of overwintering beetles in the site; predictions of beetle densities for individual trees were not reliable.     

Estimating age‐dependent survival when juveniles resemble females: Invasive ring‐necked parakeets as an example

Many species only show sexual dimorphism at the age of maturity, such that juveniles typically resemble females. Under these circumstances, estimating accurate age‐specific demographic parameters is challenging. Here, we propose a multievent model parameterization able to estimate age‐dependent survival using capture–recapture data with uncertainty in age and sex assignment of individuals. We illustrate this modeling approach with capture–recapture data from the ring‐necked parakeet Psittacula krameri. We analyzed capture, recapture, and resighting data (439 recaptures/resightings) of 156 ring‐necked parakeets tagged with neck collars in Barcelona city from 2003 to 2016 to estimate the juvenile and adult survival rate. Our models successfully estimated the survival probabilities of the different age classes considered. Survival probability was similar between adults (0.83, 95% CI = 0.77–0.87) and juveniles during their second (0.79, 95% CI = 0.58–0.87) and third winter (0.83, 95% CI = 0.65–0.88). The youngest juveniles (1st winter) showed a slightly lower survival (0.57, 95% CI = 0.37–0.79). Among adults, females showed a slightly higher survival than males (0.87, 95% CI = 0.78–0.93; and 0.80, 95% CI = 0.73–0.86, respectively). These high survival figures predict high population persistence in this species and urge management policies. The analysis also stresses the usefulness of multievent models to estimate juvenile survival when age cannot be fully ascertained.     

High water-loss rates and rapid dehydration in the burying beetle, Nicrophorus marginatus

Abstract.  During the summer months, there is a high mortality of burying beetle (Coleoptera: Silphidae) species in pitfall traps containing dry soil. The present study investigated the possibility that the burying beetle Nicrophorus marginatus is highly susceptible to death from desiccation. In the laboratory, adult beetles lose 1–5% body mass per hour in low humidity conditions (25–30% relative humidity), depending on temperature. This rate of water-loss results in a 50% mortality within 7–16 h at temperatures between 16 and 28 °C. Nicrophorus marginatus produces oral and anal defensive secretions when disturbed but these secretions do not significantly contribute to the high rate of water loss. Beetles readily drink and thus beetles with access to water or in high humidity conditions suffer near zero mortality. For comparison, the similarly sized mesic bess beetle (Coleoptera: Passalidae), Odontotaenius disjunctus , and a large nocturnal tiger beetle (Coleoptera: Cicindelidae), Amblychila cylindriformis , are exposed to dehydrating conditions. Nicrophorus marginatus have higher rates of water-loss (by 26% and 79%, respectively). It is predicted that the diurnal N. marginatus uses behavioural mechanisms to avoid lethal water loss. In the field, N. marginatus displays a strong bimodal activity pattern, avoiding the hottest hours of the day. The significance of these findings is that N. marginatus defies normal predictions of association between water-loss rates and habitat type, and has extended its range into apparently unfavourable habitats despite high water-loss rates. In addition, the results suggest that researchers should provide moisture when studying burying beetles, including the endangered American burying beetle ( Nicrophorus americanus ).     

Size-dependent perennation bud formation and its effect on seedling establishment inVeronicastrum sibiricum ssp.japonicum

Seedling establishment of a perennial,Veronicastrum sibiricum ssp.japonicum colonizing at 1,600 m altitude in Nikko National Park was studied for two years. About half of the first-year seedlings died during the growing season and 45–74 per cent of the seedlings surviving the first summer died during the following winter season. This winter mortality depended on the end-of-season size of the seedling in the previous autumn and was caused mainly by the absence of perennation buds. Seedlings which had emerged on bare ground attained larger size and had higher winter survival than seedlings in a mat of moss. Seedlings germinating in early season attained larger size and had higher probability of winter survival than seedlings germinating in later season.     

Linking summer foraging to winter survival in yellow pine chipmunks (Tamias amoenus)

This study links summer foraging and scatter-hoarding to winter larder-hoarding and winter survival in yellow pine chipmunks (Tamias amoenus) by comparing patterns of time allocation and winter larder contents in 2 years with very different levels of resource availability. In 2003, seed production and the number of trees and shrubs producing seeds were high. In 2004 seed crops were small. Chipmunks allocated more time to foraging when food resources were scarce (66% in 2004) compared to when they were abundant (39% in 2003). Increased time allocated to foraging in 2004 corresponded to significant decreases in time allocated to vigilance, resting, and social interactions. When seeds were scarce (i.e., in autumn 2004), chipmunks spent more time searching for cached food items than gathering seeds from plants or the ground surface. Despite the increase in foraging effort, the edible mass and caloric contents of larders were significantly smaller in 2004. In the year with low seed production, the diversity of seed species found in larders increased, and many of these seeds were of species that ripened in summer. When autumnal seed production by Jeffrey pine seeds was high, Jeffrey pine seeds were nearly the exclusive food item found in larders. Larder contents would have provisioned chipmunks for an estimated 116-257 days in 2003 and but only 6-111 days in 2004. It is likely that all chipmunks would have survived the winter of 2003 (duration ~110-120 days). However, none of the larders recovered in 2004 contained enough food to have provisioned the inhabitant for the ~148-158 days of winter.     

All-year reproduction and possible thermal melanism in <Emphasis Type="Italic">Amblystogenium pacificum</Emphasis> (Coleoptera: Carabidae) on the sub-Antarctic Ile de la Possession (Iles Crozet)

In the fell-field habitat at 140–270 m altitude on the sub-Antarctic Ile de la Possession, Iles Crozet, the dimorphic beetle Amblystogenium pacificum Putzeys (Coleoptera: Carabidae) was studied monthly throughout the southern year 1993–1994 by timed hand collections involving turning stones on the rocky fell-field. There were many mating couples and females containing well-developed eggs in every month and the species reproduced throughout the year, with no evidence of a winter cessation of egg production. However, the frequency of mating couples was about twice as great in the summer half-year (October–March) as in the winter half (April–September). Relatively small differences in monthly mean temperatures between the summer and winter at Iles Crozet render all-year reproduction not unexpected when comparisons are made with the temperatures and phenology of Carabidae in cool habitats in other parts of the world. Mating pairs were collected at unusually high frequencies for carabids. The frequency of such pairs, taken per unit effort by hand collecting, was highest in October–December, lower from January to May and intermediate from June to September. Comparison of beetles taken as solitary, non-mating individuals with those taken as mating couples showed that in summer, females of the black morph of this dimorphic species were more frequently involved in mating than expected, but no differences were detected in winter. Female beetles were divided into younger individuals (putative age 1–2 years) and older females (putative age 3–5 years or more) by the amount of claw-wear on individuals of this species. This division indicated that about 40% of the older females showed no egg development when captured, but only about 20% of the younger females were non-breeding. However, of the reproducing beetles, young and old females matured similar numbers of eggs. An altitude transect showed that the proportion of the black morph increased significantly with altitude. Studies at 140–270 m showed that a significantly smaller proportion of the older females of the brown morph were breeding than in the younger brown individuals, and this effect was particularly evident in the colder winter period. No such age differences existed amongst the black morph.     

Natural enemies of the convergent lady beetle, Hippodamia convergens Guérin-Méneville: Their inadvertent importation and potential significance for augmentative biological control

Each year, Hippodamia convergens adults are collected from their overwintering sites in California and made commercially available for aphid control in agriculture and in home gardens. This study examines the prevalence of natural enemies encountered in commercially available H. convergens from three commercial suppliers. Microsporidia were detected in individuals from 13 of 22 shipments (0.9% prevalence, range 0–3%). Spore dimensions (3.9 ± 0.1 × 2.5 ± 0.1 μm; n = 50) were similar to those of Nosema hippodamiae (reported previously from H. convergens), Nosema tracheophila and Nosema coccinellae (reported from other coccinellids). Because H. convergens are sold in large quantities, thousands of microsporidia-infected beetles could be released each time H. convergens are used. Three distinct eugregarines were observed in H. convergens (0.2% prevalence) but none were similar in size to Gregarina barbarara, the only eugregarine reported from H. convergens. Although eugregarines are usually considered as mutualist symbionts or weak pathogens, the relationship between eugregarines and coccinellids is not fully understood. The hymenopteran parasitoid, Dinocampus coccinellae, was found in beetles from all shipments (8% mean parasitism, range: 3–15%). Significantly more females were parasitized than males. The importation and release of D. coccinellae undermine the success of biological control programs and may artificially increase the number of parasitoids in an area where H. convergens are released. Verticillium sp. (2% prevalence; range: 0–9%) was also observed but its role is unknown. Beetle quantities and sex ratios were also assessed.     

Residency Time as an Indicator of Reproductive Restraint in Male Burying Beetles

The cost of reproduction theory posits that there are trade-offs between current and future reproduction because resources that are allocated to current offspring cannot be used for future reproductive opportunities. Two adaptive reproductive strategies have been hypothesized to offset the costs of reproduction and maximize lifetime fitness. The terminal investment hypothesis predicts that as individuals age they will allocate more resources to current reproduction as a response to decreasing residual reproductive value. The reproductive restraint hypotheses predicts that as individuals age they will allocate fewer resources to current reproduction to increase the chance of surviving for an additional reproductive opportunity. In this study, we test for adaptive responses to advancing age in male burying beetles, Nicrophorus orbicollis. Burying beetles use facultative biparental care, but the male typically abandons the brood before the female. Previous work in male burying beetles has suggested several factors to explain variation in male residency time, but no study has observed male behavior throughout their entire reproductive lifetimes to determine whether males change residency time in an adaptive way with age. We compared residency time of males that reproduced biparentally, uniparentally, and on different-sized carcasses to determine if they used an adaptive reproductive strategy. Males did not increase residency time as they aged when reproducing biparentally, but decreased residency time with age when reproducing uniparentally. A decrease in parental care with age is consistent with a reproductive restraint strategy. When female age increased over time, males did not increase their residency time to compensate for deteriorating female condition. To our knowledge, this is the first test of adaptive reproductive allocation strategies in male burying beetles.     

Developmental mortality increases sex‐ratio bias of a size‐dimorphic bark beetle

1. Given sexual size dimorphism, differential mortality owing to body size can lead to sex‐biased mortality, proximately biasing sex ratios. This mechanism may apply to mountain pine beetles, Dendroctonus ponderosae Hopkins, which typically have female‐biased adult populations (2 : 1) with females larger than males. Smaller males could be more susceptible to stresses than larger females as developing beetles overwinter and populations experience high mortality. 2. Survival of naturally‐established mountain pine beetles during the juvenile stage and the resulting adult sex ratios and body sizes (volume) were studied. Three treatments were applied to vary survival in logs cut from trees containing broods of mountain pine beetles. Logs were removed from the forest either in early winter, or in spring after overwintering below snow or after overwintering above snow. Upon removal, logs were placed at room temperature to allow beetles to complete development under similar conditions. 3. Compared with beetles from logs removed in early winter, mortality was higher and the sex ratio was more female‐biased in overwintering logs. The bias increased with overwinter mortality. However, sex ratios were female‐biased even in early winter, so additional mechanisms, other than overwintering mortality, contributed to the sex‐ratio bias. Body volume varied little relative to sex‐biased mortality, suggesting other size‐independent causes of male‐biased mortality. 4. Overwintering mortality is considered a major determinant of mountain pine beetle population dynamics. The disproportionate survival of females, who initiate colonisation of live pine trees, may affect population dynamics in ways that have not been previously considered.     

Effect of dung burial by the dung beetle Bubas bison on numbers and viability of Cryptosporidium oocysts in cattle dung

Cryptosporidium oocysts were inoculated into fresh dung (∼1.2 × 10⁴ oocysts per gram wet weight) and fed to dung beetles to assess the effect of dung burial by the dung beetle Bubas bison on the distribution of the oocysts in small cores of soil in the laboratory. The experiment consisted of five replicates of each of two treatments; controls (dung but no dung beetles) and the experimental treatment (inoculated dung and seven pairs of dung beetles). After 5 days, when approximately 90% of the dung was buried, the surface and buried dung was recovered and subsampled. The oocysts in the subsamples were recovered and enumerated using qPCR. Oocyst viability was evaluated using an assay based on the exclusion or inclusion of two fluorogenic vital dyes, 4′,6-diamidino-2-phenylindole (DAPI) and propidium iodide (PI). Results revealed that overall 13.7% of oocysts remained on the surface and 86.3% of oocysts were buried. The viability of oocysts in buried dung was only 10% compared to oocysts the surface dung (58%). Therefore, widespread dung burial by B. bison during the winter months could substantially reduce the numbers of Cryptosporidium oocysts available to be washed into waterways following winter rains.