749Carrion Beetles (Coleoptera: Silphidae) CMaculation usually various dark shades of brown or gray, with various spots or markings; some lighter or colorful. Adults are nocturnal. Larvae are borers in trunks and limbs. Host plants are recorded in a large number of plant families, espe-cially all those with larger tree species. A number of species are economic pests of forest trees.

References

Arora GS (1976) A taxonomic revision of the Indian species of the family Cossidae (Lepidoptera). Rec Zool Surv India 69:1–160

Barnes W, McDunnough JH (1911) Revision of the Cossidae of North America. Contr Nat Hist Lepidoptera North Am 1:(1)1–35, pl 1–7

Buser R, Huber W, Joos R (2000) Cossidae – Holzbohrer. In: Schmetterlinge und ihre Lebensräume: Arten-Gefährdung Pro-Schutz. Schweiz und angrenzenden Gebiete, 3:97–116, pl 2. Pro Natura-Schweizerische Bund fuer Naturschutz, Basel

Schoorl JW Jr (1990) A phylogenetic study on Cossidae (Lep-idoptera: Ditrysia) based on external adult morphology. Zoologische Verhandlingen 263:1–295, 1 pl

Seitz A (ed) (1912–1937) Familie: Cossidae. In: Die Gross-Schmetterlinge der Erde, 2:417–431, pl 52–55 (1912); 2(suppl):241–245 (1933); 287, pl 16 (1934); 6:1264–1287, pl 167, 169, 181–184 (1937); 10:807–824, pl 93, 96–99 (1933); 14:540–551, pl 79–80 (1929). A. Kernen, Stuttgart

Carposinidae

A family of moths (order Lepidoptera). They com-monly are known as fruitworm moths. Fruitworm Moths Butterflies and Moths

Carrion Beetles (Coleoptera: Silphidae)

derek s. sikesUniversity of Alaska Museum, Fairbanks, AK, USA

Despite the association of silphids with carrion, which is often repugnant to humans, the biology

of these organisms includes a rich and complex array of fascinating evolutionary and ecological phenomena.

Silphids (Fig. 26) have the largest bodies and are the most conspicuous of the staphylinoid bee-tles. However, the family is not very species rich by beetle standards, containing only 183 extant species. Commonly, they are known as “carrion beetles” due to their frequent association with vertebrate carcasses. They are sometimes referred to as “large carrion beetles” to distinguish them from other beetles associated with carrion, such as those in the family Leiodidae (which are sometimes called “small carrion beetles”). Most species eat carrion although many will also prey on carrion associated insects, such as maggots, or other carrion beetles. Some species are phytophagous, or exclusively pre-daceous, while at least one species has been found only in dung. The primary food source for the larvae of most species, however, is vertebrate car-rion. A radical departure from this ancestral life history pattern is seen in the species Nicrophorus pustulatus which, although capable of breeding on carrion, has recently been discovered to be a para-sitoid of snake eggs – perhaps the only known example of a parasite of a vertebrate that kills and consumes its host (in this case, snake embryos).

The family contains two subfamilies, each of which specializes on a different size of carrion. Carrion feeding members of the subfamily Silphi-nae, which lack parental care, prefer large verte-brate carcasses (>300 g), and are often found on megafaunal carcasses, such as elk, moose, or bison. They must share these carcasses with vertebrate scavengers and a large suite of necrophilous insects such as the larvae of blow- and fleshflies, some of which become prey for the beetles. Adults of the subfamily Nicrophorinae, which display parental care and complex subsocial nesting behaviors, generally only breed by monopolization of a small carcasses (<300 g, usually <100 g) such as those of birds or rodents. These beetles remove small carcasses from the competitive arena of flies, ants, and other scavengers by burial into a sub-terranean nest – hence their common name of

750 Carrion Beetles (Coleoptera: Silphidae)C“burying beetles” or “sexton beetles.” This remark-able behavior attracted the attention of early naturalists and remains the focus of research today.

Distinguishing Characteristics and Relationships

The family as a whole has only one, somewhat ambiguous, synapomorphy (diagnostic character) – a bulge on the posterior quarter of each elytron. However, silphids can be easily recognized by a combination of characters including their necro-philous habits (most species), their size (usually 1–2 cm), their weakly to strongly clubbed anten-nae, their very large scutellum, which is sometimes as wide as their head, and their tricostate elytra. There is strong evidence indicating silphids are closely related to members of the family Staphylin-idae. Some evidence suggests silphids may actually belong inside the family Staphylinidae (which would require changing the family Silphidae into a subfamily of the Staphylinidae).

Members of the subfamily Nicrophorinae do possess an unambiguous synapomorphy – a pair of stridulatory files on the dorsal surface of abdominal segment 5 which are used for auditory communication between adults before mating (courtship songs) and between adults and larvae during development, and for defense when dis-turbed. The files are scraped by the underside of the elytral apices when the beetles pump their abdomens forward and backward. Paired stridula-tory files are absent from the Silphinae but adults of the basal genera Ptomaphila (Australia) and Oxelytrum (Neotropics) possess stridulatory morphology – at least one species of Oxelytrum has been observed stridulating. These beetles have spines on the underside of the elytra that can be scraped by the abdominal intersegmental mem-brane when the beetles move their abdomens from side to side – a phenomenon that has yet to be studied in the silphines. Because silphines do not nest, presumably the only function of their stridu-lation would be defense.

Morphology

Adult

Length 7–45 mm (usually 12–20 mm); ovate to moderately elongate, and slightly to strongly dors-oventrally flattened (Silphinae). Frontoclypeal (epistomal) suture absent (Silphinae), or present as fine line (Nicrophorinae). Antennae are 11-seg-mented but appear as 10-segmented in Nicro-phorinae due to reduced second segment fused to third segment; ending in 3-segmented club, usually preceded by two or three enlarged but sparsely setose segments (Silphinae and basal Nicrophorinae) or antennomeres 9–11 forming a large club (Nicrophorus). Pronotum with lateral edges complete, sometimes explanate. Scutellum large – often as wide as head. Elytra truncate, exposing 1–5 abdominal tergites in Diamesus, Necrodes, and Nicrophorinae; not truncate in remaining Silphinae, covering abdomen; never striate; in Silphinae bearing 0–3 raised costae or carinae per elytron (present but indistinct in Nicrophorinae); with raised callus near posterior end of outermost costa; epipleura usually well-developed and with ridge complete almost to apex. The elytra of most Nicrophorus, Ptomascopus and Diamesus species usually have broad colored bands or spots (fascia and maculae) extending laterally to meet epipleura. Abdomen with sternite 2 not visible between hind coxae but visible later-ally of metacoxae; sternites 3–8 visible in females, 3–9 visible in males. Legs with five tarsal segments per tarsus. Males usually with broadly expanded protarsal segments and longer protarsal setae (midtarsal also expanded in male Diamesus), pro- and midtarsi of female similar.

Larvae

Length 12–40 mm, campodeiform (most Silphi-nae) or eruciform (Nicrophorinae); elongate, more or less parallel-sided to ovate, slightly to strongly flattened, relatively straight or slightly curved

751Carrion Beetles (Coleoptera: Silphidae) Cventrally. Body surfaces heavily pigmented and heavily sclerotized (Silphinae), or lightly pig-mented and lightly sclerotized (Nicrophorinae). Stemmata 6 (Silphinae) or 1 (Nicrophorinae) on each side. Mandibles lacking mola. Thoracic terga and abdominal terga and sterna consisting of one or more sclerotized plates, without patches or rows of asperities, each tergum with 1 (Silphinae) or 2 (Ptomascopus) lateral tergal processes extending beyond edges of sterna or without such processes (Nicrophorus) but with four spinose projections along posterior margin of abdominal terga. One or two segmented, well developed urogomphi.

Classification, Diversity, and Distribution

In recent years there have been some changes to the classification and newly described species added, so a current classification, with updated species counts, and distributional information is provided below (Table 7). The family currently stands at 183 species.

The distribution of an organism is the result of both ecology and evolutionary history. Silphids, especially the nicrophorines, are rare in warmer climates, such as lowland tropical forests, and vir-tually absent from dry climates like deserts – these ecological constraints certainly have limited their distribution in places like Africa, Australia, and Tibet. A few silphids in northern Africa survive in cooler, wetter mountainous regions but the Sahara presumably prevents southward dispersal. The family Silphidae is thought to have originated in the northern hemisphere on the paleocontinent of Laurasia. The subfamily Nicrophorinae best repre-sents this with only three species in territory once part of the southern landmass of Gondwana. These three species are thought to have radiated down the Andes of South America, having survived in the cooler, montane climate.

The nicrophorines are distributed in the northern hemisphere, with species radiations hav-ing occurred in the Malay Archipelago, resulting

in various endemic island species restricted to montane habitats, and into South America along the Andes. None are found in Africa south of the Sahara, in Australia (Fig. 25), or Antarctica. These beetles were once thought to be absent from the Indian subcontinent south of the Himalayas, but there may be a population of the recently described species Nicrophorus sausai in Meghalaya India, a mountainous region isolated from the Himalayas. This unusual, and perhaps relict, population begs additional study and confirmation.

The silphines are more widespread than the nicrophorines, with greater representation on Gondwanan areas. This is thought to be related to their greater generic diversity (12 genera) and possible greater age. There are four species in Australia and New Guinea (Ptomaphila, 3 endemic species; Diamesus 1 species) and a larger radia-tion in South America than is seen in the nicro-phorines (Oxelytrum, 8 species). It has been suggested that this radiation of the Silphinae into South America and consequently Australia (via Antarctica) took place 50–60 million years ago producing these, the only two silphid genera endemic to the southern Hemisphere. There are also three silphine species in South Africa (Thanatophilus, 2 species; Silpha, 1 species) and an entire silphine genus (Heterotemna, 3 species) endemic to the Canary Islands off the northwest coast of Africa. However, as with the nicro-phorines, most species of the Silphinae are found in the northern hemisphere, although they seem to be somewhat more tolerant of warm habitats than are the nicrophorines. This tolerance is per-haps due to their preference for larger carcasses which they do not (and could not) defend from competitors. Nicrophorus species, probably due to their requirement for small carcasses that can be buried and defended, do not appear to compete well with the ants, flies and carrion-associated scarab beetles that are more abundant in warmer habitats.

Together, the Silphidae show an amphitropical or amphipolar distribution, i.e. they are restricted to northern and southern temperate zones but

752 Carrion Beetles (Coleoptera: Silphidae)CCarrion Beetles (Coleoptera: Silphidae), Table 7 Carrion beetle classification, species counts, and distribution

Order Coleoptera

Superfamily Staphylinoidea

Family Silphidae Latreille, 1807 15 genera, 183 species

Subfamily Silphinae Latreille, 1807 12 genera, 111 species

Aclypea Reitter, 1884 13 species, Holarctic

Dendroxena Motschulsky, 1858 2 species, Eurasia

Diamesus Hope, 1840 2 species, Asia, Australia

Heterosilpha Portevin, 1926 2 species, West Nearctic

Heterotemna Wollaston, 1864 3 species, Africa: Canaries

Necrodes Leach, 1815 3 species, Holarctic

Necrophila Kirby and Spence, 1828 17 species, Holarctic

subgenus Necrophila Kirby & Spence, 1828

subgenus Eusilpha Semenov- Tian-Shanskij, 1890

subgenus Calosilpha Portevin, 1920

subgenus Deutosilpha Portevin, 1920

subgenus Chrysosilpha Portevin, 1921

Oiceoptoma Leach, 1815 9 species, Holarctic

Oxelytrum Gistel, 1848 8 species, SW Nearctic/Neotropical

Ptomaphila Kirby & Spence, 1828 3 species, Australia, New Guinea

Silpha Linnaeus, 1758 25 species, Eurasia, Africaa

subgenus Silpha Linnaeus, 1758

subgenus Phosphuga Leach, 1817

subgenus Ablattaria Reitter, 1884

Thanatophilus Leach, 1815 24 species, Holarctic & Africa, Madagascar

Subfamily Nicrophorinae Kirby, 1837 3 genera, 72 species

Eonecrophorus Kurosawa, 1985 1 species, Nepal

Ptomascopus Kraatz, 1876 3 species, Asia

Nicrophorus Fabricius, 1775 68 species, Holarctic, N Africa, S America, SE Asia

aOne species of Europe, Silpha trisits Illiger, has been introduced and established in North America (southern Quebec)

753Carrion Beetles (Coleoptera: Silphidae) C

generally absent from the intervening tropics (with the exception of tropical montane habitats).

The lesser generic diversity of the nicrophorines (3 genera) compared to the silphines, combined with their almost pure Laurasian distribution, sup-ports preliminary estimates for a younger age of the main radiation in the Nicrophorinae (the genus Nicrophorus) based on fossil and molecular divergence dating methods. All known fossils of the genus Nicrophorus are Eocene or younger, less than 50 million years old, with the majority being known from the Pleistocene. Molecular dating methods provide a preliminary, and wide, range for the radiation of the genus having happened 50–24 million years ago. The transition to the Oligocene from the Eocene is thought to represent the most dramatic climatic change of the Ceno-zoic era, in which the Mesozoic “hot house” world was transformed into the Neogene “ice house” world that persists today. Given the absence of Nicrophorus from lowland tropical habitats and

the preference of these organisms for cooler cli-mates, it seems reasonable to infer these beetles may have radiated during this cooling event of the Oligocene. Roughly concurrent with this cooling event, many modern rodent families appeared and radiated – and these would have been ideal prey items for Nicrophorus beetles. In addition to a small-mammal radiation during this time, most modern bird orders and families appeared between the early Eocene and the late Oligocene-early Miocene, during a period of intense diversifica-tion. Small birds are also ideal prey items for Nicrophorus.

The family is thought to have originated in the Old World, and the subfamily Nicrophorinae certainly shows this pattern in which all but one of the five genera/subgenera are endemic to Asia. The New World has the minority of world species for all groups that are also found in the Old World, and has only two endemic genera: Oxelytrum and Heterosilpha. One new species of Nicrophorus,

Carrion Beetles (Coleoptera: Silphidae), Figure 25 Map of the subfamily Nicrophorinae (Silphidae). 6,736 localities from 17,250 specimens examined showing the known distribution (99% of records are Nicrophorus). The lack of records throughout much of Russia is almost certainly a collection artifact whereas the absence of records in Australia, sub-Saharan Africa, most of India and South America is not. A corresponding map for the subfamily Silphinae has not yet been prepared – see text for description of the distribution of the Silphinae.

754 Carrion Beetles (Coleoptera: Silphidae)C

N. hispaniola, was recently discovered and described in the Dominican Republic. This was the first Nicrophorus described in the New World since 1925, bringing the total to 21 New World species of Nicrophorus. There are 25 species of Silphinae in the New World, combined with the 21 species of nicrophorines yielding 46 species of silphids in the New World.

Ecology

Silphids are most frequently encountered at verte-brate carrion but are sometimes found associated with dung or fungi, or at electric lights. Most species will prey on fly larvae or other insects pres-ent on carcasses or dung, in addition to eating the

carcass itself. Some species are phytophagous (11 species of Aclypea) while others are predacious (Dendroxena, some Silpha and possibly Ptomascopus zhangla, a poorly known and recently discovered species from China). The silphine Necrodes surina-mensis as an adult feeds primarily on fly larvae but can survive on carrion alone. The majority of silphids that have been studied are nocturnal or crepuscular (active at sundown and sunrise), which might help avoid predation by birds.

Some species of the genus Nicrophorus have become model organisms for research in ecology, physiology, and behavior – particularly dealing with questions about parental care, the evolution of sociality, competition, and other behaviors of nesting organisms (e.g., brood parasitism). There have been over 150 behavioral ecology studies on these species in the past 25 years. Subjects of these studies include, for example, the ability of adults to regulate their brood size to match the size of the carrion resource via control of the number of eggs produced and subsequent parental culling of “extra” larvae by cannibalism. Other subjects investigated include adult competition and fights to win a carcass, pheromone emission, adult strid-ulatory communication (between parents and larvae, precopulatory, and defensive), duration and explanation for paternal care, and antimicro-bial properties of anal and oral secretions, among many others. Biparental care, as seen in Nicrophorus, is rare in insects in general, and has been the focus of much investigation.

The typical progression from discovery to new offspring for Nicrophorus species proceeds as follows: A small vertebrate carcass, like that of a mouse, is found soon after nightfall and often on the day of its death. If numerous Nicrophorus beetles find the carcass the beetles begin to fight to dominate the resource. Larger bodied beetles tend to win these competitions with losers retreating, sometimes with minor injuries (missing leg parts or cuts in their wing covers). The loser females sometimes lay eggs near the carcass and some of her offspring might enter and develop in the nest of the winning beetles. The beetles’ fights usually

Carrion Beetles (Coleoptera: Silphidae), Figure 26 Nicrophorus olidus Matthews, a silphid found in Honduras and Mexico; female, dorsal and lateral view.

755Carrion Beetles (Coleoptera: Silphidae) Cresult in a single species remaining, with smaller bodied species having been excluded. Within the larger species the males fight the males and the females fight the females until the largest male and largest female remain in control of the carcass. Courtship stridulation occurs and can lead to rejection of the male by the female if he cannot stridulate as expected. This may help beetles iden-tify conspecifics but no one has carefully investi-gated how this is accomplished. The mechanisms by which closely related species avoid hybridiza-tion are equally unstudied. The male and female pair work together to bury the carcass by digging beneath it. If the substrate is too tough the pair might move the carcass to more suitable ground by laying beneath it and moving it with their legs. If a male finds a carcass and no females are present he will emit a pheromone to attract a female. Sometimes males without carcasses will emit pheromone to attract females with whom they try to mate. Females, like all insects, can store sperm for later use and sometimes can find and bury car-casses alone, using sperm from prior mating to fertilize her eggs.

It can take 5–24 h for a carcass to be secured below-ground (with smaller-bodied species tend-ing to bury less deeply than larger bodied species). The carcass is rolled into a ball that minimizes its surface area. Fur or feathers are removed and a brood chamber is built that will house the carcass and the developing larvae. The carcass is treated with oral and anal secretions that help preserve the resource from microbial decay. The female will then lay eggs based on the mass of carcass (between 10 and 50 eggs is typical) although more eggs are typically laid than larvae that will be reared. Because body size is critical to winning contests for carcasses, larger bodied offspring will be more likely to successfully reproduce than smaller bod-ied offspring. A carcass resource can yield either many small burying beetle offspring, or fewer large burying beetle offspring. This selection pressure has resulted in a behavior known as filial cannibal-ism – the parents kill and consume “extra” larvae that would otherwise lower the average body size

of the resulting offspring. These beetles, therefore, regulate the size of their brood carefully – both by laying a clutch size appropriate to the mass of the carrion resource, and by later “fine-tuning” the clutch size if too many eggs hatch by eating the late arriving larvae.

Parent beetles stay with the larvae during their approximate 2-week development period – defending them against possible usurpers (this being the major advantage of paternal care). The parents also tend the larvae, maintain the brood ball and regurgitate food for the larvae. The burying beetles are unusual among insects in having peak levels of juvenile hormone (normally considered a gonadotropic hormone in adult insects) during the early parental period when the ovaries are small. Parental regurgitations to young larvae increase larval growth rates, and in some species, are essential for molting from the first to second instar. The larvae molt between each of three instars and either pupate to adult and overwinter as adults or overwinter as a final larval instar “pre-pupa.”

This life history of Nicrophorus species is based on finding, concealing and monopolizing small carcasses before their competitors. However, they cannot exclude all interested parties – in addition to a vigilance that prevents usurpation by other Nicrophorus adults, the parents must con-tend with both bacterial and fungal decay of the carcass. Recent studies in both North America and Japan have shown that the treatment of the carcass with oral and anal secretions by Nicrophorus adults greatly reduces the microbial decay, and a number of antimicrobial agents have been identified.

Other animals that often accompany the beetles into their nest include both nematodes and mites (Acari). The nematode-beetle relationship is poorly known with considerable potential for future work. At least two nematode species, Rhab-ditis stammeri, and R. vespillonis, have been docu-mented as associates of Nicrophorus vespilloides and N. vespillo, respectively, although it is certain that many more, probably undescribed, nematode species associate with silphids. The nematodes

756 Carrion Beetles (Coleoptera: Silphidae)Cbreed on the carrion and their offspring disperse to new carrion resources with the new generation of beetles – traveling in the gut of the beetle larvae and using the adult hindgut and/or genitalia for transport to a new carcass. Nematodes have been observed in laboratory settings to reach enormous population sizes – causing the beetles to abandon the carcass – but it is unknown if this happens in the wild. The new generation of nematodes will form large aggregations on the surface of the carcass, each will arch upwards and start to wave. They will also form small, living, waving towers by climbing on one another. This behavior brings the nematodes in contact with the beetles, onto which they climb. It is not known how or if the beetles have adapted to competitive pressure from these nematodes, nor is it known to what degree the nematodes reduce the beetles’ fitness.

The mite-beetle relationship is better under-stood than that of the nematodes, but remains one of the more complex and rich areas for future research. It is not known how many species of sil-phids carry mites (phoretic associates) but most of the Nicrophorus species that have been studied ecologically carry them. Like the nematodes, the mites’ life cycle is tied to that of the beetles with the deuteronymphs (the last pre-adult stage) dis-persing phoretically on the adult beetles. Mites present on silphine species probably are using them as alternate hosts opportunistically until they can transfer to a burying beetle. Many of the mites appear to be host-specific, and considerable taxonomic work remains to done with them. Over 14 species of mites from four families (Parasitidae, Macrochelidae, Uropodidae, and Histiomatidae) were found on Nicrophorus species in Michigan, USA. The most frequently encountered and well-studied mites in this system are those in the genus Poecilochirus (Mesostigmata: Parasitidae). Initial work in the 1960s indicated an apparent mutualis-tic mite-beetle relationship resulting from the mites’ predation on fly eggs that would otherwise hatch and compete with beetle offspring. However, more thorough examination of this relationship has found much greater complexity – including

examples of mutualism, commensalism, and para-sitism, varying with species and conditions. What was once thought to be a single species of mite, Poecilochirus carabi, has since been discovered to be a species complex of several morphologically similar, but reproductively isolated species that are specific to their host beetle species. Only a few of these cryptic mite species have been described or examined in detail. It is likely that most of the 69 known Nicrophorus species have their own (prob-ably undescribed) Poecilochirus species.

An even more poorly-known symbiotic rela-tionship involving the Silphidae awaits study: nematodes of the family Allantonematidae have been reported as parasites of the burying beetles’ Poecilochirus mites!

Conservation

In the USA, much attention has been focused recently on the American burying beetle, Nicro-phorus americanus Olivier, a federally listed endan-gered species and one of five “giant” species in the genus. As recently as the 1930s, this species was considered to be common over most of the eastern half of the North American continent. However, it now occurs in <10% of its former range (popula-tions are now restricted to a few islands offshore of Rhode Island and Massachusetts and the western periphery of the historic range). This species was first listed in 1989 and represents an unusual case of species endangerment in that there are no appar-ent causal factors for its decline that simultaneously explain why the eight other co-occurring Nicro-phorus species have not declined. Many weakly supported hypotheses have been suggested, includ-ing DDT contamination, extinction of the passenger pigeon, deforestation, artificial lighting, loss of carrion availability, and an unknown, intrinsic, genetic effect. One important difference between N. americanus and its congeners is that this species requires larger carcasses (>80 g) than its congeners to maximize its reproductive success. Subsequent work and review of the literature points to a “best,”

757Carrion Beetles (Coleoptera: Silphidae) Calbeit provisional, explanation of this species’ decline based on (i) known population declines of optimally sized carrion “prey” species such as ground nesting birds and the passenger pigeon, and (ii) increased vertebrate scavenger and conge-ner competition for the reduced carrion available. The greater pressure from vertebrate scavengers may have resulted from competitive release after the loss of larger predators (such as the gray wolf [Canis lupus] and the mountain lion [Felis con-color]) and an increase in habitat fragmentation and edge habitats. Nicrophorus americanus may have declined because it is experiencing greater vertebrate and congener competition for a reduced resource base. The species is being bred in captivity and work is underway to establish new popula-tions. The attention it has received due to its federal protection has helped its prospects considerably.

Given the well-documented and recent rise in mean global temperature, some conservationists are worried about montane island endemics that cannot survive in the warmer lowlands. As our climate changes, the size of these cooler, montane habitats will contract as they gradually move higher in eleva-tion. There are at least ten Nicrophorus species endemic to the higher elevations of various islands in the Malay Archipelago. These species, among many other similarly adapted organisms, could become threatened with extinction if their cooler, montane habitats start to disappear. Some are already living at the highest elevations available to them.

Recent Research

Recent work on these beetles has resulted in some interesting discoveries. In addition to 11 newly described species since 1999, primarily from Asia, there has been phylogenetic work underway which has suggested that the relatively high species richness of the genus Nicrophorus may have resulted from a rapid radiation – a burst of evolution. This radiation was possibly coinci-dent with the global cooling during the Oligocene and the subsequent radiation of small birds and

mammals – and possibly caused by the key inno-vation of small carcass monopolization.

From other recent research we have learned that females with a carcass will not attack males who have recently been in contact with a carcass and typically cared for a brood. These males are considered to have a “breeder’s badge”, a profile of cuticular hydrocarbons that identifies them as parental – and those males lacking this scent are attacked. This addresses questions of how these social beetles recognize each other.

It had already been determined that adults cannot recognize their own larvae from those of other couples, nor even, of other species of nicrophorines (in Japan, Ptomascopus larvae are sometimes brood parasites, mixed into broods of Nicrophorus concolor larvae and raised by Nicrophorus parents). The mechanism by which parents can minimize such brood parasitism is temporal – they kill larvae that arrive too early or too late around the window of time that their own larvae appear.

Our understanding of the biology and evolu-tion of the Silphidae progresses, with continuing work on the phylogenetics, reproductive behaviors of basal lineages, brood parasitism, communal breeding, endocrinology, use of stable isotopes to determine larval diet, and host shifts, although many questions remain uninvestigated. Decomposer Insects Beetles (Coleoptera)

References

Ratcliffe BC (1996) The carrion beetles (Coleoptera: Silphidae) of Nebraska. Bull Univ Nebraska State Mus 13:1–100

Peck SB (2001) Silphidae Latreille, 1807. In: Arnett RH, Thomas MC (eds) American beetles: archostemata, myxophaga, adephaga, polyphaga: staphyliniformia, vol 1. CRC Press, Boca Raton, FL, pp 268–271

Scott MP (1998) The ecology and behavior of burying beetles. Ann Rev Entomol 43:595–618

Sikes DS (2005) Silphidae. In: Kristensen NP, Beutel RG (eds) Handbook of zoology, vol IV Arthropoda: Insecta part 38, Coleoptera, Beetles, vol I: Morphology and systematics (Archostemmata, Adephaga, Myxophaga, Polyphaga partim) (RG Beutel RAB Leschen, eds) Walter de Gruyter, Berlin, NY, pp 288–296

758 CarsidaridaeCSikes DS, Newton AF, Madge RB (2002) A catalog of the

Nicrophorinae (Coleoptera: Silphidae) of the world. Zootaxa 65:1–304

Carsidaridae

A family of bugs (order Hemiptera, superfamily Psylloidea). Bugs

Carter, Herbert James

GeorGe hanGayNarrabeen, NSW, Australia

Herbert James Carter was born on the 23rd of April 1858 in Marlborough, Wiltshire, England. He was educated in England, receiving his Bach-elor of Art Degree in Cambridge. At the age of 24 he migrated to Australia and took up the position of Mathematical Master at Sydney Grammar School. Later on, in 1902, he was appointed as Principal of Ascham Girls’ School in Sydney where he worked until his retirement in 1914. Although he was a devoted educator, his interest in entomology and his contribution to knowl-edge of the Australian insect fauna were very significant. He became interested in entomology soon after his arrival in Australia and produced his first paper on Australian Coleoptera in 1905. He traveled and collected extensively in New South Wales, Victoria, Tasmania, South Australia and Western Australia. He collaborated with A.M. Lea and a number of other entomologists of his era, including with K.G. Blair, the Coleopterist of the British Museum. During his long life he published 65 papers, including major works on Tenebrionidae, Buprestidae and Colydiidae. He described 55 genera and 1,234 species new to science. After retirement he continued his ento-mological work until his sudden death on the 16th of April 1940, in the Sydney suburb of Wahroonga.

References

Carter HJ (1933) Gulliver in the bush – wanderings of an Australian entomologist. Angus & Robertson, Sydney, Australia, 234 pp

Zimmerman EC (1993) Australian weevils, vol 3. CSIRO, East Melbourne, pp 493–494

Carthaeidae

A family of moths (order Lepidoptera) also known as Australian silkworm moths. Australian Silkworm Moths Butterflies and Moths

Carton

The paper manufactured by Hymenoptera for nest construction.

Carrier

An inert material serving to dilute a pesticide, and to carry it to its target.

Carrying Capacity

The theoretical maximum population size that an area can support indefinitely within defined set of conditions.

Casebearer Moths (Lepidoptera: Coleophoridae)

John B. heppnerFlorida State Collection of Arthropods, Gainesville, FL, USA

Casebearer moths, family Coleophoridae, com-prise over 1,525 species worldwide, with most being Palearctic (1,082 sp.) and in the genus,