Download References Carposinidae Carrion Beetles (Coleoptera: Silphidae)

Carrion Beetles (Coleoptera: Silphidae)
Maculation 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, especially 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 (Lepidoptera: Ditrysia) based on external adult morphology.
Zoologische Verhandlingen 263:1–295, 1 pl
Seitz A (ed) (1912–1937) Familie: Cossidae. In: Die GrossSchmetterlinge 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 commonly are known as fruitworm moths.
 Fruitworm Moths
 Butterflies and Moths
Carrion Beetles (Coleoptera:
Silphidae)
Derek S. Sikes
University of Alaska Museum, Fairbanks, AK,
USA
Despite the association of silphids with carrion,
which is often repugnant to humans, the biology
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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 beetles. 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 predaceous, while at least one species has been found
only in dung. The primary food source for the
larvae of most species, however, is vertebrate carrion. 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 parasitoid 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 Silphinae, which lack parental care, prefer large vertebrate 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 subterranean nest – hence their common name of
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Carrion Beetles (Coleoptera: Silphidae)
“burying beetles” or “sexton beetles.” This remarkable 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 necrophilous habits (most species), their size (usually
1–2 cm), their weakly to strongly clubbed antennae, 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 Staphylinidae. 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 disturbed. The files are scraped by the underside of
the elytral apices when the beetles pump their
abdomens forward and backward. Paired stridulatory 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 membrane 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 stridulation would be defense.
Morphology
Adult
Length 7–45 mm (usually 12–20 mm); ovate to
moderately elongate, and slightly to strongly dorsoventrally flattened (Silphinae). Frontoclypeal
(epistomal) suture absent (Silphinae), or present
as fine line (Nicrophorinae). Antennae are 11-segmented but appear as 10-segmented in Nicrophorinae 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 welldeveloped 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 laterally 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), proand midtarsi of female similar.
Larvae
Length 12–40 mm, campodeiform (most Silphinae) or eruciform (Nicrophorinae); elongate, more
or less parallel-sided to ovate, slightly to strongly
flattened, relatively straight or slightly curved
Carrion Beetles (Coleoptera: Silphidae)
ventrally. Body surfaces heavily pigmented and
heavily sclerotized (Silphinae), or lightly pigmented 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 virtually 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 represents 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 having occurred in the Malay Archipelago, resulting
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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 radiation in South America than is seen in the nicrophorines (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 nicrophorines, 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 perhaps 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
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Carrion Beetles (Coleoptera: Silphidae)
Carrion 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
One species of Europe, Silpha trisits Illiger, has been introduced and established in North America (southern Quebec)
a
Carrion Beetles (Coleoptera: Silphidae)
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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.
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, supports 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 Cenozoic 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 climates, 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 diversification. 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,
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Carrion Beetles (Coleoptera: Silphidae)
Carrion Beetles (Coleoptera: Silphidae),­
­Figure 26 Nicrophorus olidus Matthews, a silphid
found in Honduras and Mexico; female, dorsal and
lateral view.
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 vertebrate carrion but are sometimes found associated
with dung or fungi, or at electric lights. Most
species will prey on fly larvae or other insects present 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 surinamensis 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 stridulatory communication (between parents and
larvae, precopulatory, and defensive), duration
and explanation for paternal care, and antimicrobial 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)
result 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 identify conspecifics but no one has carefully investigated how this is accomplished. The mechanisms
by which closely related species avoid hybridization 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 carcasses 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 tending 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 bodied 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 cannibalism – the parents kill and consume “extra” larvae
that would otherwise lower the average body size
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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 contend 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, Rhabditis stammeri, and R. vespillonis, have been documented 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
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Carrion Beetles (Coleoptera: Silphidae)
breed 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 understood 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 silphids 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) dispersing 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 wellstudied mites in this system are those in the genus
Poecilochirus (Mesostigmata: Parasitidae). Initial
work in the 1960s indicated an apparent mutualistic 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 parasitism, 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 (probably undescribed) Poecilochirus species.
An even more poorly-known symbiotic relationship 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, Nicrophorus americanus Olivier, a federally listed endangered 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 (populations 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 apparent causal factors for its decline that simultaneously
explain why the eight other co-occurring Nicrophorus species have not declined. Many weakly
supported hypotheses have been suggested, including 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,”
Carrion Beetles (Coleoptera: Silphidae)
albeit 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 congener 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 concolor]) 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 populations. 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 elevation. 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.
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mammals – and possibly caused by the key innovation 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 evolution 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
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 coincident with the global cooling during the Oligocene
and the subsequent radiation of small birds and
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
757
758
C
Carsidaridae
Sikes 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 Hangay
Narrabeen, NSW, Australia
Herbert James Carter was born on the 23rd of
April 1858 in Marlborough, Wiltshire, England.
He was educated in England, receiving his Bachelor 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 knowledge 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 entomological 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. Heppner
Florida State Collection of Arthropods,
­Gainesville, FL, USA
Casebearer moths, family Coleophoridae, comprise over 1,525 species worldwide, with most
being Palearctic (1,082 sp.) and in the genus,