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In the lead section:

Pterostichus melanarius is a type of carabid (ground beetle) of the genus Coleoptera. It is native to Europe but is increasingly found in North America after being introduced to the region in the1920s. It is a predatory beetle that eats invertebrate pests, which makes it a valuable pest control agent in agricultural settings. Additionally, the beetle has wing dimorphism which has contributed to its increasing distribution across North America.

Description
Adults of P. melanarius grow to be 12-18 mm long. They appear black with straight striations on their elytra. They "are distinguished from other Pterostichus specis by their laterobasal carina and denticulate posterior angles of the pronotum." Adults have wing dimorphism, characterized by brachypterous beetles and macropterous beetles. Brachypterous beetles have shortened hind wings and are unable to fly. Macropterous beetles have mature hind wings and are able to fly. Eggs are translucent white immediately after oviposition, but darken before hatching. The larvae have a hard dark reddish-brown head. The appearance of pupae is not well documented.

Distribution
P. melanarius is found in a wide geographical range across Europe, the United States, and Canada. It can be found in "natural, anthropogenic, and agricultural biotopes." Beetles in similar geographical ranges tend to have similar development and life cycle stages, which differs from beetles in other geographical ranges. P. melanarius can traverse 2.5 -5m daily, which can increase during the summer in search of mates. In addition to walking, flight capabilities resulting from wing dimorphism contribute to the beetle's ability to traverse wide ranges.

In Europe, the beetle can be found in Scandinavia, Italy, Greece, Ireland, Russia, and Serbia. In North America, the beetle is found on both the west and east coasts of the continent, but has been documented to have a steady movement into the center of the US. It is important to study this varying distribution to build an understanding how P. melanarius might affect its new habitats.

Habitat
P. melanarius is hygrophilic and eurythermic, meaning it is able to tolerate wet environments with a wide ranges of temperatures. The beetle can inhabit habitats including forests, meadows, urban areas, and arable land. Researchers explain that the consequence of the variability of habitats is that "the total duration of individual development may vary even within one natural climatic zone, which may result in different variants of the life cycle in the local populations occupying different biotopes." In other words, the beetle is able to adapt stages of its life cycle to sync with its various habitat. The main conditions that would have to be met for the beetle to colonize and thrive in a habitat are temperatures suitable to larvae development and availability of food resources.

Life Cycle
Mating of P. melanarius occurs during early fall months closely followed by oviposition. To maximize probability of larvae surviving and having access to resources with needing to travel, female beetles select shaded and wet environments to lay their eggs. Females typically lay about 130 eggs at a time. All larval stages happen in the soil. Colder temperatures are necessary for the larvae to develop into adults and warmer temperatures are necessary for the transition from final larval to pupae. Based on this developmental timeline, fully-developed adults emerge during late May to June.

Behavior
P. melanarius, as larvae and adults, is an omnivorous predatory beetle that feeds on insects, invertebrates, and plants. Among a wide range of prey, the most common prey of the beetles are ants, caterpillars, mollusks, seeds, and plant tissue. Additionally, several studies have verified that the beetle's have a special preference for consuming slugs. Specifically, it has been suggested that it prefers the gray garden slug, Derocera reticulatum, which is an invasive slug species. The beetle often detects prey using olfactory clues. The behavior of the beetles is often studied using pitfall trapping, which is a type of sampling method used to explore the behavior of species like beetles. It provides insight into their spatial distribution and abundance in different habitats. The results of one study that used pitfall trapping In a study that used pitfall trapping, indicate that the beetles utilize their habitats based on their hunger level and habitat quality. So, starved beetles will be found in habitats with a wider range of prey compared to satiated beetles, meaning that foraging in their habitats is dictated by hunger level. Another study explains that starved beetles practice "conspecific odour-trail avoidance", which means that the P. melanarius beetles avoid searching for prey in locations that other individuals of P. melanarius have visited. This behavior is characteristic of Lévy-flight patterns, which are optimal for widely disbursed prey that may not always be fully consumed at one time, such as slugs.

Larvae are documented to feed on slugs and also practice cannibalism.

Competition
Competition between P. melanarius and other carabid beetles depends on the region. In Europe, laboratory observations indicate that competition exists between P. melanarius and other carabid species like, P. cristatus, and that P. melanarius dominates. To decrease competition in the wild, the two species inhabit different habitats. Interestingly, in Canada, where P. melanarius is an invasive species, there has been substantial evidence of competition between P. melanarius and other carabid beetles. P. melanarius does not appear to negatively impact the diversity or abundance of other carabid beetles. It is hypothesized that this is due to the beetles filling empty ecological niches.

Competition between P. melanarius and primary prey, slugs, is significant. Dense aggregations of P. melanarius are known to be associated with dense aggregations of slugs and the beetle is capable of reducing the distribution and abundance of slug populations. Slugs can avoid attacks by the beetle by using their slime and by strategically moving through the soil to avoid predation. Despite these tactics, the beetles are such avid predators of slugs that they are considered to be of great importance for biologically controlling slug populations, which are often harmful to agriculture.

Competition between P. melanarius and their predators is also significant. The beetles have many predators, primarily birds, but also "mice, bats, hedgehogs, shrews, frogs, toads, and occasionally moles." Additionally, adults beetles and larvae are parasitized by "mites, wasps, flies, and nematodes."

Relevance to Agriculture
Since P. melanarius inhabit arable fields, they are vulnerable to agricultural practices such as tilling fields and the use of insecticides. Tilling can be harmful to the life cycle and development of the beetles as it can occur during the fall and spring, which coincide with the oviposition and final development stages of the beetles. Tillage in the spring can disturb larvae and pupae in the soil, resulting in a decrease in the number of fully developed adults able to breed that emerge later in the fall. Some studies suggest that tillage has no effect on the beetles, but the contradictory results may be due to study methodology differences.

The use of insecticides on fields that are inhabited by P. melanarius has negative consequences for the beetle because the insecticide is harmful to the beetles and its prey. Direct exposure to insecticides reduces the beetle population and their activity levels. The beetles can also be indirectly exposed to insecticides through consumption of prey affected by insecticides. According to one study, the consumption of affect prey results in total mortality for a beetle, a risk which was only slightly reduced when consuming the affected prey several days after it was exposed to insecticide. Further, insecticides can cause mortality in the beetle's prey, reducing food availability and potentially resulting in starvation of the beetles. Since beetles that are starved spend more time foraging for food, P. melanarius has increased activity in insecticide treated fields.

An important method to decrease the negative impacts of harmful agricultural practices is to utilize cover crops and intercropping. In fields that used these agricultural control methods, which often require less tillage and chemical use, there was a greater abundance and activity levels of P. melanarius. Moreover, these habitats are ideal for female P. melanarius, which prefer to oviposit in "structurally complex environments" to provide more protection their eggs and larvae.

Wing Dimorphism
Several studies have determined that P. melanarius have had more rapid and expansive distribution in regions of Canada compared to other invasive carabid species. A possible explanation is that the macropterous,(capable of flight) beetles help establish new beetle population in favorable habitats. After establishment, the majority of those beetles are replaced by brachypterous (flightless) beetles. Both morphs are inherited according to Mendelian genetics, with the flightless morph being the dominant gene and the flight morph being the recessive gene. In the process of establishing a new beetle colony in a new habitat, the flight morph has a competitive advantage, but after the new colony is established, the flightless morph has the competitive advantage and a greater portion of the population will present in this morph.

In the lead section:

Necrodes littoralis is a type of carrion beetle of the genus Necrodes. As a carrion beetle, it feeds on decaying vertebrate remains and fly larvae (maggots). It is found in countries across Europe. Its feeding behaviors make it an important asset to forensic entomology.

It is also known as the "shore sexton beetle."

Description
Adults of N. littoralis grow to be 15 to 25 mm long and have a shiny black body. The beetles can be identified by a characteristic bump about three quarters down the length of their elytra. Beetles of the order Coleoptera, like N. littoralis, have truncated elytra, which is a hardened forewing that act as armor to protect the beetle from environmental factors and predators. The reason why some beetle species have adapted a shortened protection armor is unclear. The larvae of N. littoralis are campodeiform, meaning they have a flattened body, antennae, and have well-developed legs.

Distribution
Necrodes littoralis have a Palearctic distribution  but most reported observations are from across Europe. They have been observed in Austria, Hungary, Slovakia, Czech Republic, Belgium, France, and England. The beetles have also been observed in South Korea. The Latin prefix necr- denotes corpse and the Latin word littoralis denotes a coastal environment. The name could be misleading since the beetles are not found solely by the coast. In fact, most observations report finding the beetles in woody areas or fields.

Habitat
Adult beetles and larvae are primarily found on large carrions in the late stages of decomposition. Further, the carrions are primarily found outdoors, though there have been observations of the beetles on carrions that are indoors. Researchers hypothesize that the beetles have difficulty accessing decaying bodies indoors because they cannot easily detect openings into buildings. Researchers from Italy reported the presence of N. littoralis on a human corpse in Italy for the first time in 2021. The decomposition conditions of the corpse at the time of discovery align with the consensus that the beetles inhabit corpses at later stages of decomposition. The corpse was found indoors, which the researchers explain that "the access to the building through the open door and the state of total neglect of the area where the corpse was found ... may have favored the indoor colonization by N. littoralis."

Life Cycle
Mating of adult beetles often happens at night. Following mating, the female lays eggs in the ground near the carrion. Although there is variation in the exact number, females lay between 50 to 70 at a time. Researchers who studied the instar development of N. littorialis explain that the beetles have three larval stages. Another study reveals that the three larval stages are first instar, second instar, and third instar. Postfeeding larva, nymph and imago are parts of the developmental stage of larvae into adult beetles. The first instar larvae are creamy white when they hatch and shift toward the carrion for food. First instar larvae are the most vulnerable. The second and third instar larvae are also creamy white after ecdysis, which is the process of insects shedding its exoskeleton. As part of the transition from third star larvae to postfeeding larva, the third star larvae burrow into the ground and form pupal chambers "thrashing the abdomen and thus compacting the soil around them. They go through the prepupal, pupal, and teneral adult stages inside the chambers." The beetles emerge from the pupal chamber after "they become fully sclerotized and colored."

Larvae Behavior:
The larvae of N. littoralis aggregate, meaning they form large and orderly groups of larva on the carrion. In these large groups, the larvae feed on the carrion. Researchers who were interested in exploring why the beetle larvae group for feeding performed experiments to test the "importance of thermal cues and ground-deposited chemical cues for the aggregation behaviour..." The experiment involved field data and lab tests. The field data consisted of analyzing the larvae growth results of previous experiments with pig carcasses. The lab tests involved collecting adult beetles, allowing 1 male to mate with 1 female, allowing the larvae to grow, and placing them on a sample carcass set-up to observe aggregation behavior. The results of the study highlight the importance of stable thermal conditions for the larval aggregations of N. littoralis as the larvae formed aggregations around a heat source of the carrion. If the heat source moved, the larvae aggregations followed by disassembling and forming a new aggregation around the new heat source. Stable thermal conditions are important for the development of the larvae into adult beetles. Notably, larvae in the later stages of development, specifically third instar larva, prefer to aggregate in cooler temperatures. The authors hypothesize that this preference allows the larvae to grow larger, though this growth happens more slowly. The third instar larva phase begins the transition to the post-feeding phase, which is when the largest larvae were observed in another study. Additionally, the aggregations form around parts of the carrion that are favorable for feeding. The data from the experiments did not support the ground-deposited chemical cues as an important motivation for aggregation behavior.

Adult Behavior:
Necrodes littoralis and other necrophages are attracted to carrions by cadaveric volatile organic compounds (VOCs), which are gases emitted into the air by the carrion. Minimal information is known about the VOCs of cadavers at late stages of decomposition that attract beetles like N. littoralis. To find out about the VOCs that attract N. littoralis, researchers tested the response of the beetles to different VOCs. The VOCs tested were "benzyl butyrate, butran-1-ol, butyric acid, cadaverine, DMDS, DMTS, indole, phenol, putrescine and skatole..." The list of VOCs tested includes VOCs released in the late decomposition stages to account for the beetle's timing of habitation. The study found that there was not a positive and significant attraction of N. littoralis to the listed VOCs. The main limitation of the study is the low quantity of VOCs and replicates used in the experiment. Ultimately, this study did not resolve the mystery of which VOCs N. littoralis are attracted to. However, the researchers did observe that adult beetles began to inhabit the carrion around the time the carrion started bloating. Bloating of a carrion result from accumulation of gasses released by decomposition. The researchers explain that future studies should focus on identifying the gasses that cause bloating to narrow down the possible VOCs that attract N. littoralis to carrions.

Adult N. littoralis are most active during the spring and summer months. Adult beetles on carrions feed on the decaying tissues, but mostly consume Diptera larvae, which is the larvae of flies, especially blow flies.

Adult N. littoralis are hypothesized to perform indirect parental care for their larvae by spreading "anal exudates" on the carrion, which produces heat on the carrion and helps direct the location of larvae aggregation to an area with suitable temperatures.

Competition:
N. littoralis are not the only insects that inhabit large carrions. The other most common inhabitant of carrions is flies, specifically blow flies (Calliphoridae). Researchers understood that both beetles and blowflies help decompose bodies but were interested in the competitive aspect of the N. littoralis and blow fly interactions. Blow flies inhabit the carrion soon after death whereas the beetles inhabit the carrion much later after death, closer to the time of the body bloating. Despite these timing differences, the two insects share similarities in their interactions with the carrion. For example, larvae of both insects form aggregations while they feed on the carrion. The authors explain that "similarities in carrion utilization prompted us to hypothesize that blow flies and Necrodes beetles compete over large carrion." The authors hypothesized that the blow flies would have access to the best territory on the carrion since they arrive first and the beetles would choose the remaining available areas of the carrion. This hypothesis was tested by evaluating experiment results of a previous study involving pig carcasses. Another hypothesis proposed by the authors was that the beetles "compete with blow flies by killing the larvae that are prior on in their peak feeding phase...These predictions were tested in behavioral laboratory assays." The most important result of the study is that the blow flies and the beetles have a competitive relationship. This is made evident by the observation that flies reduce the available area of feeding on the carrion for the beetles, which is an indirect effect on the beetles, and that the beetles directly affect the flies by consuming the fly larvae. Interestingly, the beetles kill the fly larvae that are the youngest and smallest, indicating that consuming fly larvae is not a strategy to eat more food, but a strategy to reduce the competition over the carrion.

Relevance to Forensic Entomolgy:
Necrodes littoralis are a useful tool for forensic entomologists as analysis of their behaviors and growth can reveal evidence in death cases. Post-mortem interval (PMI) is a measurement used to identify how long a body has been dead. PMI can be determined by creating a developmental model of the activity of carrion beetles, like N. littoralis. Researchers conducted a study comparing individual rearing vs. aggregation rearing of N. littoralis to develop a standardized approach to PMI estimation. The results of the study indicate that beetles reared indivudally have a higher mortality, take more time to develop and are smaller in size. On the other hand, beetles reared in aggregation have lower mortality, take less time to develop, and are larger in size. The authors suggest that for forensic entomologists to get more precise PMI results from modeling the beetle's activities, it is best to create a model that rears the beetles in aggregation.

Catering was originally done by servants of wealthy elites. Butlers and house slaves, which were often black, were in a good position to become caterers. Essentially, caterers in the 1860s were "public butlers" as they organized and executed the food aspect of a social gathering. A public butler was a butler working for several households. Bogle took on the role of public butler and took advantage of the food service market in the hospitality field. Caterers like Bogle were involved with events likely to be catered today, such as weddings and funerals. Bogle also is credited with creating the Guild of Caterers and helping train other black caterers. This is important because catering provided not only jobs to black people but also opportunities to connect with elite members of Philadelphia society. Unfortunately, the clientele of caterers became the middle class, who could not afford lavish gatherings and increasing competition from white caterers led to a decline in black catering businesses.