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''Nicrophorus vespilloides'' is a
burying beetle Burying beetles or sexton beetles, genus ''Nicrophorus'', are the best-known members of the family Silphidae (carrion beetles). Most of these beetles are black with red markings on the Elytron, elytra (forewings). Burying beetles are true to th ...
described by
Johann Friedrich Wilhelm Herbst Johann Friedrich Wilhelm Herbst (1 November 1743 – 5 November 1807) was a German natural history, naturalist and entomologist from Petershagen, Minden-Ravensberg. He served as a chaplain in the Prussian army. His marriage in Berlin, 1770, with E ...
in 1783. The beetles vary widely in size and can present with a range of anywhere from 12 mm to 20 mm in size. They have two conspicuous orange-yellow bands on the elytra. The color of the antennae are an important distinguishing feature, being totally black. The color of their orange and black markings is multifunctional, as they are conspicuous to avian predators. In general, they present a unique ecological niche, which is their evolution of aposematism, or the strategy they use to warn predators through their conspicuous signals. The wing cases of these beetles possess a squarish shape and are notably shorter in length than their abdomens, indicating a distinct physical characteristic of the species. Overall, there are reported differences in coloration and chemical defenses among the sexes. This is hypothesized to stem from conflicting selection pressures for the different sexes. They also produce anal fluid or exudate from their abdomen when they feel threatened. These anal fluids further contribute to the antimicrobial defense of these beetles.


Geography

These insect species boast a wide distribution that spans an extensive range of geographic areas, including the far northern regions of
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
within
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, to
Siberia Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
, and reaching into various parts of
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, notably
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
and
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
. Additionally, their geographic range stretches into
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, covering northern parts of the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
and including much of southern
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
. This broad distribution underscores the beetles' exceptional adaptability and resilience, enabling them to flourish in a diverse array of environmental contexts. They are particularly prevalent in both the
Palaearctic The Palearctic or Palaearctic is a biogeographic realm of the Earth, the largest of eight. Confined almost entirely to the Eastern Hemisphere, it stretches across Europe and Asia, north of the foothills of the Himalayas, and North Africa. Th ...
and
Nearctic The Nearctic realm is one of the eight biogeographic realms constituting the Earth's land surface. The Nearctic realm covers most of North America, including Greenland, Central Florida, and the highlands of Mexico. The parts of North America ...
biogeographic realms, which is indicative of their ability to adapt and survive under a wide spectrum of environmental conditions. Within the expansive Palaeearctic region, these beetles occupy a wide ecological niche, making their home in environments ranging from the low-lying plains to the higher elevations of alpine regions. They can be found in various habitats that notably encompass dense woodlands, open heathlands, landscaped parklands, and even residential gardens, showcasing their remarkable versatility. In contrast, within the Nearctic region, their habitat preferences appear to be more selective. Here, they are primarily found in specific types of environments, such as sphagnum bogs, marshy areas, and the fringes of upland regions. This suggests a more specialized adaptive nature to particular environmental conditions prevalent in these parts of North America. This extensive and varied geographical distribution, along with their diverse habitat preferences, highlight the beetles' wide-ranging adaptability and the ecological resilience they exhibit across different regions of the world.


Description

The beetles of ''N. vespilloides'' have highly variable body size that ranges from 12–20 mm. They have two conspicuous orange-yellow bands on the elytra. The color of their antennae are completely black. Their orange and black markings serve as a warning sign to avian predators that defends them from attack. Their distinct wing cases are squarish in shape and are shorter than their abdomens.


Activity

The adult beetles are known to follow specific seasonal patterns of activity, which are intricately linked to the cyclical changes in their natural habitats and the current climate that they live in. Typically, these insects emerge from dormancy and become notably active during the spring months, with their activity beginning around spring in early April and May. This period marks the beginning of their most active phase, which continues through the months of spring and summer, extending into the late autumn. Within this active phase, there are particularly significant peaks in activity observed in May, a time when environmental conditions are most favorable for their survival and for engaging in reproductive behaviors. Following this peak, there is a notable resurgence in activity during the late summer months, suggesting a bi-modal pattern of activity. This pattern is not arbitrary but rather is a strategic adaptation to the environmental conditions that are most conducive to their survival and the successful reproduction and nurturing of their offspring.


Bi-parental care

An aspect of the behavioral ecology of these beetles is their commitment to biparental care, where both male and female parents are deeply involved in the care and nurturing of their offspring. This biparental care strategy is characterized by a thoughtful and balanced investment of resources not only towards activities directly related to reproduction, such as egg-laying and the guarding of larvae but also towards maintaining a robust immune function. This dual focus highlights a highly evolved and sophisticated approach to ensuring the survival and well-being of their offspring while, at the same time, preserving the health and vitality of the adult beetles. This approach to parental care is particularly noteworthy as it indicates a level of complex social behavior and cooperation between male and female beetles, contributing significantly to their success as a species. By investing in both their immediate reproductive success and their longer-term health and survival, these beetles demonstrate an adaptive strategy that allows them to thrive across a diverse range of environmental landscapes, underlining the complexity and sophistication of their life strategies in facing the challenges of their ecosystems.


Burying behavior

''Nicrophorus vespilloides'' shares its reproductive behavior with a sister species ''
Nicrophorus nepalensis ''Nicrophorus nepalensis'' ( Chinese: 尼泊爾埋葬蟲 or 橙斑埋葬蟲), commonly known as burying beetle, is widespread across tropical and subtropical countries in Asia. It belongs to the order Coleoptera and the family Silphidae, and is p ...
''. Both species locate vertebrate carcasses and bury them underground to use them as a food source for their offspring. While their respective habitats and ecological niches certainly change the exact mechanism of burying tendencies, both species engage in the general framework of this form of bi-parental care.


Sister beetle

What had been considered ''Nicrophorus vespilloides'' in mid and eastern Canada and northeastern USA was determined by Sikes et al. in 2016 to be a separate, overlooked sister species of ''Nicrophorus vespilloides'' that had been named by Kirby in 1837. This sister species, ''Nicrophorus hebes'' Kirby, is restricted to ''Sphagnum'' bogs and
marsh In ecology, a marsh is a wetland that is dominated by herbaceous plants rather than by woody plants.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p More in genera ...
es. ''Nicrophorus vespilloides'' occurs throughout the northern Palearctic, Alaska and northwestern Canada where it is found in open
forest A forest is an ecosystem characterized by a dense ecological community, community of trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, ...
habitats. The restriction of its sister species ''N. hebes'' to bogs in North America has been attributed to competition with its closely related congener, '' N. defodiens'' which in this area is found in forest habitats. ''N. hebes'' reproduces exclusively in bogs in North America and is never found in adjacent (<) forested habitat in the
Mer Bleue Mer Bleue Bog is a protected area in Gloucester, Ontario, an eastern suburb of Ottawa in Eastern Ontario, Canada. Its main feature is a sphagnum bog that is situated in an ancient channel of the Ottawa River and is a remarkable Boreal ecosystem ...
bog area near
Ottawa Ottawa is the capital city of Canada. It is located in the southern Ontario, southern portion of the province of Ontario, at the confluence of the Ottawa River and the Rideau River. Ottawa borders Gatineau, Gatineau, Quebec, and forms the cor ...
, Ontario, Canada.


Mites

There are also a number of
phoretic Phoresis or phoresy is a temporary Commensalism, commensalistic relationship when an organism (a phoront or phoretic) attaches itself to a host organism solely for travel. It has been seen in tick, ticks and mite, mites since the 18th century, ...
(hitch-hiking)
mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...
s that are associated with ''N. vespilloides''. These include '' Pelzneria'' nr. ''crenulata'', ''
Macrocheles merderius ''Macrocheles'' is a genus of mites in the family Macrochelidae that includes over 80 described species. Species These 81 species belong to the genus ''Macrocheles'': * '' Macrocheles analis'' (Hyatt & Emberson) * '' Macrocheles baliensis'' ...
'', and '' Uroobovella'' nr. ''novasimilis'' and the largest mite ''Poecilochirus carabi''. ''P. carabi'' is not attached by any physical means (such as a secreted anal stalk in the case of ''M. merderius'') to ''N. vespilloides''. When the males or females of ''N. vespilloides'' have finished breeding on a carcass the
deutonymph Mites start as an egg and then may pass through up to six instars: prelarva, larva, protonymph, deutonymph, tritonymph, and adult. These developmental stages may look different or may be omitted depending on the mite group. All mites have an adult ...
s of ''P. carabi'' roam freely about the body of the beetles as they search for new carcasses to reproduce. It had been proposed that ''P. carabi'' deutonymphs, on arrival at a new carcass dismounted from the beetles and consumed fly eggs and larvae which would have competed for the beetle larvae for food. This relationship which benefited the beetles has been described as mutualistic. However, it has been shown that adults of ''P. carabi'' consume the eggs of ''N. vespilloides'' and that this has direct and negative effects on the reproduction of this beetle species. This is one of the most well studied of the burying beetles with over 1,000 citations found via Google Scholar. ''N. vespilloides'' is also used as a model organism in the study of social immunity.


Reproduction and parental care

The reproductive success of burying beetles is intricately linked to their unique ecological niche, specifically their reliance on securing a small vertebrate carcass for breeding purposes. The discovery of such a carcass is a pivotal moment in the beetles' reproductive cycle, triggering a significant increase in juvenile hormone (JH) levels. It's noteworthy that in other species, elevated JH levels have been associated with the suppression of the immune system, hinting at a possible trade-off in burying beetles where there might be a down regulation of the immune response during the critical period of breeding. This adaptation underscores the beetles' prioritization of reproduction, even at the potential cost of reduced immune defenses. Upon locating a suitable carcass, the beetle parents engage in a collaborative effort to bury it beneath the soil. This is a meticulous preparation process where any fur or feathers are carefully removed, and the carcass is shaped into a compact ball. This ball is then coated with a concoction of antimicrobials and secretions, a deliberate action taken by the beetles to slow down the decomposition process. It is in the vicinity of this carefully prepared carcass, now nestled within the soil, that the beetles choose to lay their eggs. A few days after the eggs are laid, the larvae hatch and instinctively navigate their way to the carcass, which serves as both their food source and nursery. The adult beetles then take on the role of providers, offering their young pre-digested food and protecting them from potential predators and competitors. This nurturing phase is critical, and should one parent abandon the effort or reduce their participation, the remaining partner compensates by ramping up their caregiving efforts through the efforts of practicing bipartialism. Instances of both parents deserting their offspring are exceptionally rare in this species, underscoring the high degree of parental investment typical of burying beetles. Moreover, the dynamic of parental decision-making in this context is fascinating, suggesting a complex interplay of negotiation and cooperation between the male and female. This is not a scenario where decisions are made unilaterally; rather, it involves continuous interaction and mutual adjustment between the parents, pointing to a sophisticated level of social coordination and communication. The decision-making process is likely a result of a negotiation involving repeated interactions between the male and female, rather than a single decision made by each parent independently. The link between parental care and the health and growth of the offspring is profound and well-documented. Research indicates that the quality of food provisioning by the parents, particularly in terms of the carcass's freshness, plays a significant role in determining offspring health and development. For instance, one study highlighted that offspring tended to fare better in terms of growth and health when their parents were breeding on higher quality, less decomposed carcasses. This contrasted with scenarios where the parents were utilizing lower quality, more decomposed food sources, thereby underscoring the critical role of parental effort and resource quality in the developmental success of the burying beetles' young. This intricate balance between resource acquisition, parental care, and offspring development highlights the complexity of the burying beetles' reproductive and social behavior, offering valuable insights into the evolutionary pressures and adaptations that shape their life history strategies.


Immunity

In the world of burying beetles, the balance between the imperatives of reproduction and the maintenance of immune function unfolds as a complex natural behavior for these beetles further showcasing the remarkable capabilities of both male and females for their practice of biparental care. This intricate dynamic involves a meticulous calibration where the energy and resources dedicated to reproduction must be judiciously balanced against those reserved for immune defense mechanisms. Central to the beetle's immune arsenal is the enzyme
Phenoloxidase Polyphenol oxidase (PPO; also polyphenol oxidase i, chloroplastic), an enzyme involved in fruit browning, is a tetramer that contains four atoms of copper per molecule. PPO may accept monophenols and/or ''o''-diphenols as substrates. The ...
(PO), a critical player in the cellular immune response and vital for the cuticle's hardening process. During the reproductive phase, there's a noticeable increase in PO activity, suggesting a potential impact on parental investment strategies by directly influencing reproductive success metrics. This link is pivotal because the quality and extent of parental care are directly correlated with the growth and survival rates of the offspring, making the number and mass of offspring reliable barometers of parental effort. The life cycle of burying beetles offers a vivid illustration of this equilibrium, with juvenile hormone levels experiencing a surge as larvae emerge—a phase coinciding with a noted decrease in PO levels. However, research indicates that PO levels can be upregulated in response to injury, even as larvae partake in feeding on the carcass. This aspect is particularly pertinent in scenarios involving encapsulation challenges, where a pronounced increase in PO activity would be imperative. Such an adaptive immune response to physical injury underscores the beetles' ability to navigate the potential drawbacks of immunosuppression, effectively masking any trade-offs between immune functionality and reproductive endeavors. This nuanced dynamic underscores a sophisticated strategy employed by the beetles, deftly allocating resources to ensure the sustenance and protection of their progeny while concurrently safeguarding their survival against the myriad environmental challenges they face. This complex interplay between reproductive strategies and immune function not only highlights the evolutionary adaptiveness of burying beetles but also offers a fascinating glimpse into the resource management strategies critical for their survival and reproductive success.


Genes

In the context of burying beetles, which predominantly feed on the carrion of vertebrates, an intriguing aspect of their biology is their genetic composition, particularly in relation to their digestive capabilities. These beetles possess a very limited set of genes related to the breakdown of
lignocellulose Lignocellulose refers to plant dry matter (biomass), so called lignocellulosic biomass. It is the most abundantly available raw material on the Earth for the production of Biofuel, biofuels. It is composed of two kinds of carbohydrate polymers, c ...
, a primary component of plant cell walls. Specifically, their genome includes just a single gene from the
glycoside hydrolase family 9 In molecular biology, glycoside hydrolase family 9 is a family of glycoside hydrolases. Glycoside hydrolases are a widespread group of enzymes that hydrolyse the glycosidic bond between two or more carbohydrates, or between a carbohydrate and a ...
(GH9) and a few from the glycoside hydrolase family 1 (GH1). This genetic makeup is somewhat unexpected, given that their diet does not typically include lignocellulosic materials, which are abundant in plant matter. Despite the absence of a lignocellulose-rich diet, there exists a complex and highly integrated interaction between the metabolic processes of the beetles and their gut microbiome. This synergistic relationship plays a crucial role in the beetles' ability to digest their food efficiently. Moreover, the gut microbiome assists in the detoxification processes, which is vital for the beetles' ability to consume carrion, a food source that might otherwise be toxic or harmful due to decomposition and the presence of potentially pathogenic microorganisms. Furthermore, the collaboration between the beetles and their gut
microbiota Microbiota are the range of microorganisms that may be commensal, mutualistic, or pathogenic found in and on all multicellular organisms, including plants. Microbiota include bacteria, archaea, protists, fungi, and viruses, and have been found ...
extends to the protection of their food source. The microbial community within the beetles' gut contributes to warding off spoilage and inhibiting the growth of competing microorganisms. This ensures that the carrion remains a viable food source for the beetles and their offspring for a longer period. This interaction highlights the importance of the gut microbiome in supplementing the beetles' digestive capabilities, allowing them to extract necessary nutrients from their specialized diet and protect their niche from competitors. This complex interplay between the beetles' metabolic functions and their gut microbiome underscores the adaptive strategies these organisms employ to thrive in their ecological niche, despite the seemingly limited genetic toolkit for digesting their primary food source.


Future research

This is one of the most well-studied of the burying beetles with over 1,000 citations found via Google Scholar. ''N. vespilloides'' is also used as a model organism in the study of social immunity. In future research, this beetle will continue to be a model used to understand the many complex social behaviors that exist such as parental care, mating conflict, sibling-sibling conflict, and the genetic architecture of these behaviors.


See also

Nicrophorus quadripunctuatus


References

{{Taxonbar, from=Q577020 Silphidae Beetles of North America Beetles of Europe Beetles of Asia Beetles described in 1783 Taxa named by Johann Friedrich Wilhelm Herbst