Odontotaenius Disjunctus Research Articles

Wood fibers are a crucial microhabitat for cellulose- and xylan- degrading bacteria in the hindgut of the wood-feeding beetle Odontotaenius disjunctus.

Wood digestion in insects relies on the maintenance of a mosaic of numerous microhabitats, each colonized by distinct microbiomes. Understanding the division of digestive labor between these microhabitats- is central to understanding the physiology and evolution of symbiotic wood digestion. A microhabitat that has emerged to be of direct relevance to the process of lignocellulose digestion is the surface of ingested plant material. Wood particles in the guts of some termites are colonized by a specialized bacterial fiber-digesting microbiome, but whether this represents a widespread strategy among insect lineages that have independently evolved wood-feeding remains an open question. In this study, we investigated the bacterial communities specifically associated with wood fibers in the gut of the passalid beetle Odontotaenius disjunctus. We developed a Percoll-based centrifugation method to isolate and enrich the wood particles from the anterior hindgut, allowing us to access the wood fibers and their associated microbiome. We then performed assays of enzyme activity and used short-read and long-read amplicon sequencing of the 16S rRNA gene to identify the composition of the fiber-associated microbiome. Our assays demonstrated that the anterior hindgut, which houses a majority of the bacterial load, is an important site for lignocellulose digestion. Wood particles enriched from the anterior hindgut contribute to a large proportion of the total enzyme activity. The sequencing revealed that O. disjunctus, like termites, harbors a distinct fiber-associated microbiome, but notably, its community is enriched in insect-specific groups of Lactococcus and Turicibacter. Our study underscores the importance of microhabitats in fostering the complex symbiotic relationships between wood-feeding insects and their microbiomes. The discovery of distinct fiber-digesting symbionts in O. disjunctus, compared to termites, highlights the diverse evolutionary paths insects have taken to adapt to a challenging diet.

Sex-specific effects of a parasite on stress-induced freezing behavior in a natural beetle-nematode system.

Some animals react to predation threats or other stressors by adopting a freezing posture in an attempt to avoid detection, and the duration of this behavior usually corresponds with individual personality, such that timid individuals freeze longer. Despite decades of research on this or related behaviors (thanatosis), never has the impact of parasitism been considered. Parasites could prolong the duration, if hosts are less motivated to move (i.e. lethargic), or they could reduce it, if hosts are motivated to forage more to compensate for energy drain. We examined this behavior within a natural beetle-nematode system, where hosts (horned passalus beetles, Odontotaenius disjunctus) are parasitized by a nematode, Chondronema passali. We exposed beetles (n = 238) to four stressors in our lab, including noise, vibration, light and inversion, and recorded how long they adopt a frozen stance. Afterward, we determined nematode burdens, which can range from dozens to hundreds of worms. Beetles tended to freeze for 20 seconds on average, with some variation between stressors. We detected no effect of beetle mass on the duration of freezing, and this behavior did not differ in beetles collected during the breeding or non-breeding season. There was a surprising sex-based difference in the impact of nematodes; unparasitized females remained frozen twice as long as unparasitized males, but for beetles with heavy nematode burdens, the opposite was true. From this we infer that heavily parasitized females are more bold, while males with heavy burdens would be more timid. The explanation for this finding remains elusive, though we can rule out many possibilities based on prior work on this host-parasite system.

Characterization of Cross-Species Transmission of Drosophila melanogaster Nora Virus.

Drosophila melanogaster Nora virus (DmNV) is a novel picorna-like virus first characterized in 2006. Since then, Nora virus has been detected in several non-Drosophila species, including insects in the Orders Hymenoptera, Lepidoptera, Coleoptera, and Orthoptera. The objective of this study was to determine if DmNV could infect individuals of other species of invertebrates besides D. melanogaster. The presence of DmNV in native invertebrates and commercially available stocks was determined. Laboratory-reared D. yakuba, D. mercatorum, Gryllodes sigillatus, Tenebrio molitor, Galleria mellonella, and Musca domestica were intentionally infected with DmNV. In addition, native invertebrates were collected and D. melanogaster stocks were purchased and screened for DmNV presence using reverse transcription-polymerase chain reaction (RT-PCR) before being intentionally infected for study. All Drosophila species and other invertebrates, except M. domestica, that were intentionally infected with DmNV ended up scoring positive for the virus via RT-PCR. DmNV infection was also detected in three native invertebrates (Spilosoma virginica, Diplopoda, and Odontotaenius disjunctus) and all commercially available stocks tested. These findings suggest that DmNV readily infects individuals of other species of invertebrates, while also appearing to be an endemic virus in both wild and laboratory D. melanogaster populations. The detection of DmNV in commercially available stocks presents a cautionary message for scientists using these stocks in studies of virology and immunology.

Variation in Larval Thermal Tolerance of Three Saproxylic Beetle Species.

Temperature is a key abiotic condition that limits the distributions of organisms, and forest insects are particularly sensitive to thermal extremes. Whereas winged adult insects generally are able to escape unfavorable temperatures, other less-vagile insects (e.g., larvae) must withstand local microclimatic conditions to survive. Here, we measured the thermal tolerance of the larvae of three saproxylic beetle species that are common inhabitants of coarse woody debris (CWD) in temperate forests of eastern North America: Lucanus elaphus Fabricius (Lucanidae), Dendroides canadensis Latreille (Pyrochroidae), and Odontotaenius disjunctus Illiger (Passalidae). We determined how their critical thermal maxima (CTmax) vary with body size (mass), and measured the thermal profiles of CWD representing the range of microhabitats occupied by these species. Average CTmax differed among the three species and increased with mass intraspecifically. However, mass was not a good predictor of thermal tolerance among species. Temperature ramp rate and time in captivity also influenced larval CTmax, but only for D. canadensis and L. elaphus respectively. Heating profiles within relatively dry CWD sometimes exceeded the CTmax of the beetle larvae, and deeper portions of CWD were generally cooler. Interspecific differences in CTmax were not fully explained by microhabitat association, but the results suggest that the distribution of some species within a forest can be affected by local thermal extremes. Understanding the responses of saproxylic beetle larvae to warming habitats will help predict shifts in community structure and ecosystem functioning in light of climate change and increasing habitat fragmentation.

Multiple lineages of Streptomyces produce antimicrobials within passalid beetle galleries across eastern North America.

Some insects form symbioses in which actinomycetes provide defense against pathogens by making antimicrobials. The range of chemical strategies employed across these associations, and how these strategies relate to insect lifestyle, remains underexplored. We assessed subsocial passalid beetles of the species Odontotaenius disjunctus, and their frass (fecal material), which is an important food resource within their galleries, as a model insect/actinomycete system. Through chemical and phylogenetic analyses, we found that O. disjunctus frass collected across eastern North America harbored multiple lineages of Streptomyces and diverse antimicrobials. Metabolites detected in frass displayed synergistic and antagonistic inhibition of a fungal entomopathogen, Metarhizium anisopliae, and multiple streptomycete isolates inhibited this pathogen when co-cultivated directly in frass. These findings support a model in which the lifestyle of O. disjunctus accommodates multiple Streptomyces lineages in their frass, resulting in a rich repertoire of antimicrobials that likely insulates their galleries against pathogenic invasion.

Walking while Parasitized: Effects of a Naturally-Occurring Nematode on Locomotor Activity of Horned Passalus Beetles

Internal parasites typically are associated with a range of negative effects on their hosts, including reduced energy, which can manifest in behavioral alterations. With this in mind, we examined effects of a naturally-occurring nematode parasite, Chondronema passali, on locomotor activity level in horned passalus beetles, Odontotaenius disjunctus from Georgia, USA. This parasite is not well-studied but can number in the thousands in severely parasitized hosts. Prior study in our lab revealed that parasitized beetles actually consume more wood than unparasitized ones do, leading us to ask here, if parasitized beetles are also more physically active. Beetles were collected from nearby forests and housed individually in our lab. We created a simple tabletop arena to observe beetle locomotor activity, which was gridded and included small stones and paper objects. We allowed individual beetles to traverse the arena for 5 min and recorded the number of grid squares crossed. Then, beetles were dissected to determine parasite presence and level of infection (on a categorical scale). A total of 140 beetles were examined across three collections. Statistical analyses of locomotor activity revealed parasite severity predicted locomotor activity, but paradoxically, lightly-infected beetles were twice as active as those without this nematode. Activity diminished with increasing worm burdens thereafter, but even the group with the most severe burdens did not move less than those with no worms. From these results we conclude that this parasite does not result in overall reduction in activity, but rather it appears to come with heightened locomotion. Alternatively, this result could stem from the fact that more active beetles are simply more likely to contract the parasite.

A novel d-xylose isomerase from the gut of the wood feeding beetle Odontotaenius disjunctus efficiently expressed in Saccharomyces cerevisiae

Carbohydrate rich substrates such as lignocellulosic hydrolysates remain one of the primary sources of potentially renewable fuel and bulk chemicals. The pentose sugar d-xylose is often present in significant amounts along with hexoses. Saccharomyces cerevisiae can acquire the ability to metabolize d-xylose through expression of heterologous d-xylose isomerase (XI). This enzyme is notoriously difficult to express in S. cerevisiae and only fourteen XIs have been reported to be active so far. We cloned a new d-xylose isomerase derived from microorganisms in the gut of the wood-feeding beetle Odontotaenius disjunctus. Although somewhat homologous to the XI from Piromyces sp. E2, the new gene was identified as bacterial in origin and the host as a Parabacteroides sp. Expression of the new XI in S. cerevisiae resulted in faster aerobic growth than the XI from Piromyces on d-xylose media. The d-xylose isomerization rate conferred by the new XI was also 72% higher, while absolute xylitol production was identical in both strains. Interestingly, increasing concentrations of xylitol (up to 8 g L−1) appeared not to inhibit d-xylose consumption. The newly described XI displayed 2.6 times higher specific activity, 37% lower KM for d-xylose, and exhibited higher activity over a broader temperature range, retaining 51% of maximal activity at 30 °C compared with only 29% activity for the Piromyces XI.

Phylogeography and demographic expansion in the widely distributed horned passalus beetle, Odontotaenius disjunctus (coleoptera: Passalidae)

Dynamic climatic oscillations during the Pleistocene had profound effects on the distribution of species across North America. Although the role of historical climate change on speciation remains controversial, the impact on genetic variation within species has been well documented. Analyses of mtDNA sequences from the cytochrome oxidase I gene (911 bp) for 115 individuals of Odontotaenius disjunctus was combined with ecological niche modelling (ENM) to infer the demographic and population differentiation scenarios under present and past conditions. We inferred three lineages that diverged during the Pleistocene and replace each other geographically across the eastern United States. One of these lineages traverses previously identified genetic barriers for terrestrial animals including the Mississippi and Apalachicola Rivers and the Appalachian Mountains. We observed overlapping ranges between two haplotype groups as well as a region of secondary contact associated with ecological transition zone in northern Florida. The two continental lineages depict a genetic signature of a recent population increase associated with expanding niche envelope, whereas the clade restricted to peninsular Florida shows stable populations in a shrinking niche envelope. Given the lack of ecological separation, overlapping distribution of haplogroups and the presence of secondary contact zones, the taxonomic status of these lineages must await robust testing using multilocus DNA data to assess species boundaries.

KEANEKARAGAMAN MAKROFAUNA TANAH DI PERKEBUNAN KOPI DESA BATU KALUNG KECAMATAN MUARA KEMUMU KABUPATEN KEPAHIANG

This study aims to determine the diversity of existing soil macrofauna at coffee plantations located at Batu Kalung village, Muara Kemumu subdistrict of Kepahiang regency. This research was conducted in February 2018 at the coffee plantations area. The methods used in this research were pitfall-trap and hand sorting. The result of the study at the site found there are 9 orders of macraofauna consisting of 11 families, 13 species, and 188 spesimen.The species found were Heterometrus lepturus, Scolopendrae morsitans, Trigoniulus corallines, Neocurtilla hexadactylla, Gryllus assimilis, Ceuthophilus maculates, Dolichoderus bituberculatus, Componotus carnelus, Blatta germanica, Blattella orientalis, Odontotaenius disjunctus, Ponthoscolex corethurus, and Hirudo medicinalis. The calculation of soil macrofauna index in coffee plantations found a number of 0.942. This indicates that that the level of soil macrofauna diversity at the coffee plantations is low. The highest soil macrofauna diversity index was found on Ponthoscolex corethurus, which is 0.146. In the contrary, the lowest diversity index was found onHeterometrus lepturus species, 0.011. In addition, The average physics-chemical factor of soil at the coffee plantation was obtained that the soil temperature is 280C, soil pH of 5.8, and soil moisture of 67

Healing while parasitized: impact of a naturally‐occurring nematode during energy‐intensive wound‐healing in a beetle

AbstractThe rigours of the daily lives of insects sometimes lead to minor injuries and wounds, which must be healed to avoid entry of pathogens and to resume normal function. Such healing requires energy, which must be diverted from other bodily reserves. What happens if energy reserves are already low, as would occur in individuals coping with internal parasites? This question is addressed in the presemt study, using horned passalus beetles (Odontotaenius disjunctus) and their naturally‐occurring nematode Chondronema passali. Oxygen consumption rates are tested at rest, as well as after an experimental wound is applied, to evaluate energy requirements of wound‐healing in parasitized and nonparasitized hosts. Furthermore, wound‐healing rates are visually tracked with a numerical scoring system to directly measure the cost of parasitism on healing. At rest, parasitized beetles show no elevation in respiration (oxygen consumption). After wounding, the oxygen consumption of parasitized beetles is 10% higher than that in nonparasitized beetles. Beetles with moderate‐ to heavy worm burdens have slower healing than those with few or no nematodes. These results show that this parasite carries little cost to the host during day‐to‐day activities, whereas, during times of immediate energy demand, there is a cost; hosts require more energy to repair wounds, and the wounds take longer to close. This conclusion leads to the question of whether this parasite is truly benign, and how many other apparently benign parasites, in insects or other animals, have similar ‘hidden’ effects.

Two Divergent Genetic Lineages within the Horned Passalus Beetle, Odontotaenius disjunctus (Coleoptera: Passalidae): An Emerging Model for Insect Behavior, Physiology, and Microbiome Research.

The horned passalus (Odontotaenius disjunctus) is one of the most extensively studied saproxylic beetles in the eastern United States. For several decades this species has been the subject of investigations into the behaviors associated with subsociality as well as physiological responses to stress, and, most recently, the composition of its gut microbiome has been closely examined. However, no published study to date has characterized this beetle’s broad-scale population genetic structure. Here, we conducted intensive geographic sampling throughout the southern Appalachian Mountains and surrounding areas and then assessed mitochondrial DNA (mtDNA) sequence variation among individuals. Unexpectedly, we discovered two divergent, yet broadly sympatric, mtDNA clades. Indeed, the magnitude of divergence between- vs. within-clades ranged from 5.9 to 7.5×, depending on the dataset under consideration, and members of the two lineages were often syntopic (i.e., found in the same rotting log). Given the potential implications for past and future studies on behavior, physiology, and the gut microbiome, we developed a simple cost-efficient molecular assay (i.e., polymerase chain reaction restriction fragment length polymorphism; PCR-RFLP) to rapidly determine mtDNA clade membership of O. disjunctus individuals. We suggest that the evolutionary processes that gave rise to the emergence and persistence of divergent sympatric lineages reported here warrant investigation, as this type of spatial-genetic pattern appears to be rare among southern Appalachian forest invertebrates.

Direct measurement of fight or flight behavior in a beetle reveals individual variation and the influence of parasitism.

How and to what degree an animal deals with potential threats is a fascinating topic that has been well-researched, particularly in insects, though usually not with the impact of parasites in mind. A growing body of work is showing how even benign parasites can affect, positively or negatively, their hosts’ physiological or behavioral reaction to threats. With this in mind we conducted an experiment using horned passalus beetles, Odontotaenius disjunctus that were naturally parasitized with a nematode Chondronema passali; we subjected beetles to simulated attacks (resembling rival fighting or predator attacks) and from videos of the encounters we quantified a suite of behaviors (antennae movement, aggressive posturing, threat displays, etc.), plus rates of alarm calls (stridulations) which all correspond to the “fight or flight” reaction. We obtained behavioral and parasite data from 140 beetles from two field collections, of which half had been housed in our lab for three weeks in conditions that would be stressful (little cover for burrowing). We observed a wide range of behaviors during the simulated attack procedure, from beetles offering little resistance to those which were extremely aggressive, though most beetles showed a moderate reaction. Alarm calling rates also varied, but surprisingly, these were not correlated with the magnitude of behavioral reactions. Also surprising was that stressful housing did not heighten the physical resistance during attacks, but did elevate alarm calling rate. Importantly, parasitized beetles had significantly reduced physical reactions to attack than those without nematodes (meaning their resistance to the attack was muted). The results concerning parasitism, coupled with prior work in our lab, indicate that the C. passali nematode depresses the hosts’ acute stress, or fight or flight, reaction (likely from its energetic cost), which may make hosts more susceptible to the very dangers that they are coping with during the stress events.

The sicker the better: nematode-infected passalus beetles provide enhanced ecosystem services.

There is growing appreciation for the role that parasites have in ecosystems and food webs, though the possibility that they could improve an ecosystem service has never been considered. In forest ecosystems, fallen trees naturally decay over time and slowly return their nutrients to the soil. Beetles in the family Passalidae play a key role by excavating tunnels and consuming wood from these logs, thereby breaking down the wood into smaller debris. In the eastern United States, the horned passalus ( Odontotaenius disjunctus) is host to a naturally occurring nematode, Chondronema passali, which appears to cause little harm to the beetles. We suspected this was due to compensatory food consumption by parasitized individuals, which we tested here. We collected and housed 113 adult beetles in individual containers with wood for three months, then determined the amount of wood each beetle had processed into fine debris and frass. We then assessed beetles for C. passali and compared wood processing rates between parasitized and non-parasitized groups. Results showed the average daily processing rate of parasitized beetles ([Formula: see text] = 0.77 g d-1) was 15% greater than that of unparasitized ones ([Formula: see text] = 0.67 g d-1). Parasitized beetles were 6% larger, and this may explain some of this pattern, though the effect of parasitism was still significant in our analysis. By extrapolating the daily rates, we estimate that 10 adult beetles without nematodes would break down approximately 2.4 kg of wood in a single year, while a group of 10 parasitized beetles would break down 2.8 kg. While our data are consistent with the idea of compensatory feeding, because these results are based on natural infections, we cannot rule out the possibility that beetles with heightened wood consumption are simply more likely to acquire the parasite. At an ecosystem level, it may not matter which is the case; parasitized beetles provide a more effective ecosystem service.

Gut anatomical properties and microbial functional assembly promote lignocellulose deconstruction and colony subsistence of a wood-feeding beetle.

Beneficial microbial associations enhance the fitness of most living organisms, and wood-feeding insects offer some of the most striking examples of this. Odontotaenius disjunctus is a wood-feeding beetle that possesses a digestive tract with four main compartments, each of which contains well-differentiated microbial populations, suggesting that anatomical properties and separation of these compartments may enhance energy extraction from woody biomass. Here, using integrated chemical analyses, we demonstrate that lignocellulose deconstruction and fermentation occur sequentially across compartments, and that selection for microbial groups and their metabolic pathways is facilitated by gut anatomical features. Metaproteogenomics showed that higher oxygen concentration in the midgut drives lignocellulose depolymerization, while a thicker gut wall in the anterior hindgut reduces oxygen diffusion and favours hydrogen accumulation, facilitating fermentation, homoacetogenesis and nitrogen fixation. We demonstrate that depolymerization continues in the posterior hindgut, and that the beetle excretes an energy- and nutrient-rich product on which its offspring subsist and develop. Our results show that the establishment of beneficial microbial partners within a host requires both the acquisition of the microorganisms and the formation of specific habitats within the host to promote key microbial metabolic functions. Together, gut anatomical properties and microbial functional assembly enable lignocellulose deconstruction and colony subsistence on an extremely nutrient-poor diet.

Pinworms (Nematoda: Thelastomatoidea) from Insects Collected in Mississippi, USA, with description of a new species of Protrellus Cobb, 1920 from the Cockroach Ischnoptera deropeltiformis Brunner von Wattenwyl, 1865 (Blattaria: Ectobiidae).

Thelastomatoid nematodes from the hindguts of several species of insects were collected in 4 localities in Mississippi, U.S.A. Protrellus browni n. sp. is described from Ischnoptera deropeltiformis (Blattaria: Ectobiidae) using light and scanning electron microscopy. This species is distinguished on the basis of its large size as well as its large eggs having one broad end tapering toward the opposite pole and with the presence of small spines on part of the shell. Protrellus aurifluus (Chitwood, 1932) Basir, 1956 was found in Parcoblatta fulvescens (Blattaria: Ectobiidae). Four other species of blattarians sampled were negative for pinworms. Additional nematodes recovered from other hosts included Zonothrix columbianus Adamson Buck 1989 (Thelastomatoidea: Pseudonymidae) from Tropisternus blatchleyi blatchleyi and Tropisternus collaris (Coleoptera: Hydrophilidae), Hystrignathus rigidus Leidy, 1850 and Xyo pseudohystrix Travassos Kloss, 1958 (Thelastomatoidea: Hystrignathidae) from Odontotaenius disjunctus (Coleoptera: Passalidae). These findings include several new host and geographical records.

The Heartrate Reaction to Acute Stress in Horned Passalus Beetles (Odontotaenius disjunctus) is Negatively Affected by a Naturally-Occurring Nematode Parasite.

There are many events in the lives of insects where rapid, effective stress reactions are needed, including fighting conspecifics to defend territories, evading predators, and responding to wounds. A key element of the stress reaction is elevation of heartrate (HR), for enhancing distribution of blood (hemolymph) to body compartments. We conducted two experiments designed to improve understanding of the insect stress reaction and how it is influenced by parasitism in a common beetle species (Odontotaenius disjunctus). By non-destructively observing heartbeat frequency before, during and after applying a stressor (physical restraint) for 10 min, we sought to determine: (1) the exact timing of the cardiac stress reaction; (2) the magnitude of heartrate elevation during stress; and (3) if the physiological response is affected by a naturally-occurring nematode parasite, Chondronema passali. Restraint caused a dramatic increase in heartrate, though not immediately; maximum HR was reached after approximately 8 min. Average heartrate went from 65.5 beats/min to a maximum of 81.5 (24.5% increase) in adults raised in the lab (n = 19). Using wild-caught adults (n = 77), average heartrates went from 54.9 beats/min to 74.2 (35.5% increase). When restraint was removed, HR declined after ~5 min, and reached baseline 50 min later. The nematode parasite did not affect baseline heartrates in either experiment, but in one, it retarded the heartrate elevation during stress, and in the other, it reduced the overall magnitude of the elevation. While we acknowledge that our results are based on comparisons of beetles with naturally-occurring parasite infections, these results indicate this parasite causes a modest reduction in host cardiac output during acute stress conditions.

Parasite Manipulation of Its Host's Physiological Reaction to Acute Stress: Experimental Results from a Natural Beetle-Nematode System.

All animals, whether vertebrate or invertebrate, must be capable of reacting to acute stressors, such as escaping from predators, and most do so with a suite of transient physiological changes that temporarily enhance survival. Some of these changes include mobilization of immune cells and increased cardiac output. A small but growing number of studies have begun to show that certain parasites appear capable of modifying such responses. We addressed this topic using a natural host and parasite system, that is, a nematode (Chondronema passali) that parasitizes horned passalus beetles, Odontotaenius disjunctus (family Passalidae), of the eastern United States. With a series of experiments, we sought to determine whether this parasite affects (1) the immune reaction to stress, (2) the output of stress-induced alarm calls, or (3) the increase in heart rate that occurs in response to acute stressors, with the stressors being mechanical or thermal. Results showed that hemocyte density increased after both stressors in nonparasitized beetles but did not increase in parasitized beetles. While mobilization of immune cells would enhance host immunity during stress, this would also be damaging to the nematode, so this scenario appears to benefit the parasite. We found no evidence that the nematode suppresses the overall reaction to stress (or prevents stress from occurring), since parasitized beetles did not differ from nonparasitized ones in alarm call rates or in heart beat frequency after exposure to mechanical stressors. Suppression of the host's normal immune reaction to stressful stimuli could translate to delayed or even reduced wound healing or pathogen resistance during these events. This project is a rare demonstration of parasite manipulation of host immune response to acute stress and should stimulate further investigations into the interactive nature of stress and parasites.

High rates of extra‐pair paternity in a socially monogamous beetle with biparental care

1. Nest construction and paternity assurance are predicted to favour biparental care in insects. The horned passalus (Odontotaenius disjunctus) is a socially monogamous beetle with biparental care that breeds in decaying logs. The genetic mating system of the horned passalus was investigated to determine if paternity assurance is likely to drive the evolution or maintenance of paternal care in this system. Parental time budgets were also examined to better understand the types and frequencies of behaviours performed by parents.2. Genotyping‐by‐sequencing revealed high levels of extra‐pair paternity, with 54.8% of offspring sired by extra‐pair males and 70% of nests containing extra‐pair young.3. More heterozygous social males were cuckolded less than more homozygous social males. Extra‐pair mating, however, seems unlikely to increase offspring genetic diversity as extra‐pair offspring were not more heterozygous than within‐pair offspring, and average brood heterozygosity did not increase with higher rates of extra‐pair paternity.4. Behavioural observations demonstrated that parents spent on average 46.5% of their time processing the decaying wood resource for larval offspring. Because resource processing is a by‐product of feeding and provides shareable benefits for all larvae in the brood, this form of paternal care could be favoured despite low paternity.

Observations ofSteinernemaNematode and Tachinid Fly Parasites in Horned Passalus Beetles,Odontotaenius disjunctus, from Georgia, U.S.A.

ABSTRACT: This report describes a series of observations made on 2 parasite species infecting a collection of horned passalus beetles (Odontotaenius disjunctus, n = 135), from Georgia, U.S.A., that were collected as larvae in the wild and were reared to metamorphosis in captivity. Tachinid fly maggots emerged from 15 beetle larvae (11%) and, when they completed their development into adults, we identified them as Zelia vertebrata, a known but little-studied parasitoid of O. disjunctus. We also observed infections by Steinernema nematodes (n = 11 cases, or 8%), which killed and consumed the carcasses and which produced thousands of infective juveniles in the rearing containers. Based on morphological measurements of infective juvenile and adult worms, these nematodes were consistent with Steinernema carpocapsae, members of which infest the surface of soils and which infect a wide range of insects. This report is the first to describe Steinernema infections in O. disjunctus, and our observations of Z. vert...

Effects of the Naturally Occurring Parasitic NematodeChondronema passaliLeidy on Lifting Strength and Captivity-Related Body Mass Patterns in the Horned Passalus Beetle,Odontotaenius disjunctus(Illiger) (Coleoptera: Passalidae)

Abstract Parasites are ubiquitous among insects and well-studied, but knowledge of their full range of host effects is not complete, especially concerning their impact on physical performance of hosts. The horned passalus beetle, Odontotaenius disjunctus (Illiger), in the USA is frequently parasitized by Chondronema passali Leidy, a nematode that can number in the thousands within the hemocoel. We performed an experiment that examined the effects of this parasite on a key performance metric, lifting strength, which should be important during fighting. We assessed beetle lifting strength using a vertically-mounted force gauge, as well as their body mass. At the end of the experiment, we dissected the beetles to determine parasite status and gender. The average beetle (1–2 g each) lifted between 300 and 400 g during the trials. Though the final sample size of non-parasitized beetles was small (n = 8), our results showed parasitized beetles tended to lift 17% less weight than non-parasitized beetles, and thi...