Encyclopedia of Entomology

6762 Pages·2008·203.16 MB·English

by  George Hangay (auth.)

D
Dactyl Dahlbom, Anders Gustav
A tarsal segment, after the first segment, that Anders Dahlbom was born in Ostgothland,
is expanded and finger-like in appearance. The Sweden, on March 3, 1806. He studied in Lund
dactyls on the front legs of mole crickets and earned a doctorate in philosophy in 1829.
(Orthoptera: Gryllotalpidae) are an adaptation Then he became an instructor in natural history
that facilitates the insect burrowing, mole-like, and preparator in the zoological museum in Lund
though the soil. in 1830. Three promotions made him professor of
 Mole Crickets and their Biological Control entomology in Lund in 1858. He explored his
 Grasshoppers, Katydids and Crickets native Sweden to collect zoological specimens
and also traveled abroad to Germany and Denmark,
and there visited museums. He married Wilhelmine
Krey, and they had six children. His area of
research was the taxonomy of Hymenoptera.
Dactylopiidae
His first two publications appeared in 1829, and
were monographs on pompilid wasps and chry-
A family of insects in the superfamily Coccoidae
sidid wasps. For almost 30 years he maintained an
(order Hemiptera). They sometimes are called
output of books and papers on Hymenoptera
cochineal insects.
while maintaining correspondence with entomol-
 Bugs
ogists in his own country and in Britain, France,
and Germany. He died in Lund on May 6, 1859.
Reference
Daddy Long-Legs
Anon (1859) Nekrolog Entomol Ztg (Stettin) 20:337–340
This term is applied to several arthropods, princi-
pally craneflies (Diptera: Tipulidae), spider-like
harvestmen (Arachnida: Opiliones or Phalang- Dalceridae
ida), and several true spiders in the family Pholci-
dae (Arachnida: Aranea), but most often Pholcus A family of moths (order Lepidoptera). They com-
phalangioides. What they all have in common is monly are known as tropical slug caterpillar moths.
that they have long, slender legs; otherwise, they  Tropical Slug Caterpillar Moths
are not closely related.  Butterflies and Moths
D
1146 Dampwood Termites
Dampwood Termites Order: Diptera
Superfamily: Empidoidea (exclusive of Dolichopo-
A group of termites in the families Kalotermitidae didae)
and Rhinotermitidae that are known to attack Family: Empididae
moist dead wood. Subfamily: Empidinae
 Termites Subfamily: Hemerodromiinae
Subfamily: Clinocerinae
Family: Atelestidae
Damsel Bugs
Subfamily: Nemedininae
Subfamily: Atelestinae
Members of the family Nabidae (order Family: Hybotidae
Hemiptera). Subfamily: Trichininae
 Bugs Subfamily: Ocydromiinae
Subfamily: Oedaleinae
Subfamily: Tachydromiinae
Damselflies
Subfamily: Hybotinae
Family: Brachystomatidae
Certain members (suborder Zygoptera) of an Subfamily: Trichopezinae
order of insects (order Odonata). Subfamily: Ceratomerinae
 Dragonflies and Damselflies Subfamily: Brachystomatinae
Together these families include nearly 4,500
described species worldwide and comprise a
Dance Flies, Balloon Flies,
morphologically diverse array of taxa, although
Predaceous Flies (Diptera:
Homalocnemis, Iteaphila, Anthepiscopus and
Empidoidea, Exclusive of
Oreogeton are still unplaced to family within this
Dolichopodidae)
classification. The vast majority of species are
predators as adults with the few exceptions being
jeffrey m. cumming1, bradley j. sinclair2 obligate flower-feeding groups that consume pol-
1Agriculture and Agri-Food Canada, Ottawa, ON, len as their only protein source. They are found in
Canada a variety of forested and open habitats where they
2Canadian Food Inspection Agency, Ottawa, ON, breed in moist soils, decaying wood, dung, and in
Canada aquatic habitats. All known larvae appear to be
predators on invertebrates.
The dance flies, balloon flies and other preda-
ceous flies that have traditionally been placed in
Distinguishing Characters
the family Empididae are now classified in four
families of Empidoidea along with the long-
legged flies of the family Dolichopodidae. With Most adult Empidoidea (exclusive of Dolichopo-
approximately 11,400 described species and didae) can be easily differentiated from the very
many more undescribed species, the Empidoidea similar Dolichopodidae by their lack of green
are one of the largest superfamilies of Diptera metallic body coloration. However a few non-
and the most diverse lineage of predaceous flies. dolichopodid Empidoidea (e.g., most Lamprempis)
The four families that were formerly classified as are metallic green and some dolichopodids are
the single family Empididae are listed below with not metallic. However, all Dolichopodidae have
their included subfamilies. the Rs wing vein originating at or near the level of
D
Dance Flies, Balloon Flies, Predaceous Flies (Diptera: Empidoidea, Exclusive of Dolichopodidae) 1147
crossvein h and have crossvein r-m situated in the a one- segmented palpus, and a labrum that is usu-
basal fourth of the wing, unlike other Empidoidea ally armed at the apex with paired epipharyngeal
that have the Rs vein originating well distal to the blades. The chaetotaxy (bristling) of the head is
level of crossvein h and have crossvein r-m distal limited, and usually includes a pair of ocellar bris-
to the basal fourth of the wing. In addition, male tles, postoculars, and sometimes additional verti-
dolichopodids have their terminalia rotated for- cal bristles.
ward beneath the preceding segments of the abdo- The thorax is generally rectangular in dorsal
men, unlike other empidoids. Some bee flies outline and is extended anteriorly in some groups.
(Bombyliidae) and asiloid groups (e.g., therevids The mesonotum is nearly flat to greatly arched
sensu lato) can also be confused with non- and dome-like. The chaetotaxy of the thorax
dolichopodid Empidoidea, but these flies gener- includes notopleurals and scutellars, and various
ally have a larger anal (cup) cell that reaches, or other dorsal bristles depending upon the group,
nearly reaches, the margin of the wing and they and the acrostichal and dorsocentral setae tend
lack any predaceous modifications on either the to be differentiated. The wings are of varied shape
legs or mouthparts. A few platypezid cyclor- and size, sometimes broadened, sometimes nar-
rhaphan flies (e.g., Microsania) can also be con- rowed, rarely reduced, with the alula and anal
fused with non-dolichopodid Empidoidea, but lobe often lacking. The wing color varies from
their arista is comprised of three articles rather clear to darkened, and is seldom patterned with
than two, and the acrostichal setae are uniserial markings. The wing venation is exceedingly var-
rather than biserial or absent. ied from relatively complete to markedly reduced.
The anal cell is closed and never reaches the wing
margin. The legs are varied in length, thickness
Morphology – Adults
and armature, and often at least one pair bears
raptorial modifications. The legs are sometimes
Adult Empidoidea (exclusive of Dolichopodidae) sexually dimorphic, with rows of pinnate bristles
are small to medium-sized flies (1–12 mm) that in females and clasping or glandular structures
are darkish to light in color and are rarely metallic in males. The empodium between the claws is
green (Fig. 1). The head is variously shaped, but is usually setiform, but is pulvilliform in some
not large and is usually narrower than the thorax. aquatic groups.
The compound eyes are generally large, with the The abdomen is subcylindrical and usually
males often holoptic (i.e., eyes meeting above in elongate, with at least some tergites bearing
the middle of the head) and the females dichoptic abdominal plaques. The male terminalia are either
(i.e., eyes separated). However in some groups unrotated or twisted 45–90° to the right. The male
the males are dichoptic, and in one major lineage genital capsule is symmetric or asymmetric and
(the Hybotinae) the females are also holoptic like has a lever-like ejaculatory apodeme that usually
the males. The antenna consists of a scape, pedi- articulates ventrally against the base of the phal-
cel, postpedicel (1st flagellomere) and a stylus. lus. The female terminalia are with or without
The pedicel is usually without the thumb-like spine-bearing acanthophorites and there is one
conus seen in most cyclorrhaphan flies that inserts spermatheca internally.
into the postpedicel. The postpedicel appears as a The family Empididae (Fig. 1a) includes small
single article that is variously shaped. The stylus is to relatively large flies (2–12 mm in length) that
usually comprised of two articles, is short to elon- usually have fairly complete wing venation, with
gate, and is either apical or more dorsally situ- vein R often forked and cell dm usually present.
4+5
ated on the postpedicel, such that it appears The prosternum is enlarged to form a precoxal
arista-like. The proboscis is short to elongate, with bridge and the legs are usually strong, long and
D
1148 Dance Flies, Balloon Flies, Predaceous Flies (Diptera: Empidoidea, Exclusive of Dolichopodidae)
Dance Flies, Balloon Flies, Predaceous Flies (Diptera: Empidoidea, Exclusive of Dolichopodidae), Figure 1
Representative Empidoidea (a) Empis aerobatica (Empididae); left – male, right – female, center – balloon
with prey (a midge) attached, USA (photograph by Eric Fisher); (b) Acarteroptera licina
(Atelestidae); female feeding on flower, Chile (photograph by Stephen Marshall); (c) Platypalpus
holosericus (Hybotidae), female, Canada (photograph by Stephen Marshall); (d) Ceratomerus sp. (virgatus
group) (Brachystomatidae), female, New Zealand (photograph by Stephen Marshall).
slender without a fore tibial gland. The male ter- The family Hybotidae (Fig. 1c) includes very
minalia are unrotated and the cerci are often small to medium-sized flies (1–9 mm) that gener-
enlarged and developed for clasping. The female ally have reduced venation, with vein R
4+5
abdomen is telescopic and pointed apically, and unbranched and cell dm often absent. Usually at
lacks acanthophorite spines on tergite 10. least one set of legs (fore, mid, or hind depending
The Atelestidae (Fig. 1b) are small flies (2–4 on the group) is raptorial and a fore tibial gland is
mm) with venation characterized by an unbranched present. The male terminalia are normally twisted
vein R and a relatively long anal cell. They lack 45–90° to the right and are often asymmetric. The
4+5
predaceous modifications on the mouthparts (i.e., female abdomen is telescopic and lacks acan-
epiphyngeal blades) and the legs are not raptorial. thophorite spines on tergite 10.
The fore tibial gland is also lacking. The male The Brachystomatidae (Fig. 1d) are small to
terminalia are unrotated and the gonocoxal apo- relatively large flies (2–11 mm) that tend to have
demes are distinctively elongated. The female fairly complete wing venation, with vein R often
4+5
abdomen is telescopic and lacks tergite 10. forked and cell dm always present. The legs are
D
Dance Flies, Balloon Flies, Predaceous Flies (Diptera: Empidoidea, Exclusive of Dolichopodidae) 1149
usually strong, long and slender and lack a fore spines along the dorsal surface of each abdominal
tibial gland. The male terminalia are unrotated segment with the terminal segment ending in a
(asymmetrically twisted to the left in some Tricho- pair of strong hooks.
pezinae), and the ejaculatory apodeme is plate-like
and narrowly fused to the base of the phallus. The
Ecology
female abdomen is truncate, and usually bears a
fringe of setae along the posterior margin of tergite
7 as well as acanthophorite spines on tergite 10. The Empidoidea (exclusive of Dolichopodidae)
represent a large group of predaceous flies, includ-
ing a few flower-visiting species. They are often
Morphology – Immatures
found in various forest habitats, on leaves, tree
trunks, damp depressions and aquatic vegetation,
The immature stages of all Empidoidea are poorly and are a dominant component of running water
known, but larvae are generally maggot-like with habitats such as streams and seeps. However, some
reduced head sclerites and a unique mandible taxa are associated with more open areas such as
made up of four to six components. Non- agricultural fields, grasslands, marshes, coastal
dolichopodid empidoid immatures are known for zones and beaches. Many groups exploit a variety
relatively few groups, but for those that are known of habitats for completion of their life cycles, using
the larvae possess either an amphipneustic, metap- different sites for larval development, swarming
neustic, or apneustic respiratory system depend- and mating, and adult feeding. Adults capture
ing on the taxon. Prolegs are generally present on various arthropod prey, including small to medium-
some abdominal segments of aquatic or semi- sized Diptera, Hemiptera, Thysanoptera, Lepi-
aquatic groups, whereas creeping welts tend to be doptera, Trichoptera, Hymenoptera, Neuroptera,
present on the abdominal segments of more ter- Ephemeroptera, Plecoptera, Coleoptera, Collem-
restrial taxa. bola, and Acari. Because of their vast species
Unlike Dolichopodidae, most other empidoid diversity and wide habitat range, predaceous
larvae do not appear to spin a cocoon before empidoid flies are important natural and poten-
pupating. There is a single report of a cocoon hav- tial biological control agents of various pest
ing been spun by a larva of a species of Drapetis insects.
(Tachydromiinae). However, no cocoon was spun A number of adult non-dolichopodid Empi-
by a larva in the related genus Megagrapha that doidea also visit flowers (Fig. 1b), presumably in
was observed pupating, suggesting that cocoon most cases to obtain nectar, but at least a few
production is not common to all Tachydromiinae groups (e.g., Iteaphila, Anthepiscopus, Anthalia,
as has been speculated. Allanthalia and Euthyneura) obtain all their pro-
Non-dolichopodid empidoid pupae vary con- tein requirements by feeding on pollen.
siderably depending on the genus and the habitat Larvae are generally found in damp soil, rot-
they occupy. Some genera (Drapetis, Megagrapha) ten wood, dung, running water, seeps and tidal
possess a pair of lengthened anterior respiratory zones, depending on the group. They appear to be
organs that arise from the anterior dorsal surface predaceous on various arthropods, particularly
of the thorax, whereas others (Hemerodromia, other Diptera larvae.
Chelifera) possess lengthened lateral abdominal As a group, Empidoidea have the potential to
respiratory processes; Neoplasta has both types of be useful bioindicators for conservation pro-
lengthened respiratory processes. Several aquatic grams and for site quality assessment. Fifteen
genera (Oreogeton, Hemerodromia, Clinocera, species of non-dolichopodid Empidoidea have
Roederiodes, Wiedemannia) have relatively stout been listed in the Red Data Book as critically
D
1150 Dance Flies, Balloon Flies, Predaceous Flies (Diptera: Empidoidea, Exclusive of Dolichopodidae)
Diversity, Evolution and
endangered, endangered or vulnerable in Great
Distribution
Britain, whereas 33 species have been listed for
Northern Belgium. Due to their predominantly
predaceous habits, they are normally not consid- The Empidoidea (exclusive of Dolichopodidae)
ered to be threatened unless their breeding are worldwide in distribution and comprise
biotopes or microhabitats are endangered. approximately 4,500 species classified in over 160
genera. In addition, numerous fossil genera and
species have been described from several Meso-
Mating Behavior
zoic and Tertiary deposits, particularly in amber
inclusions from Lebanon, Spain, New Jersey,
Many species of non-dolichopodid Empidoidea Myanmar (Burma), Siberia, Canada, the Baltic
mate on the ground or on vegetation (e.g., Region, Dominican Republic and Mexico. In
Hemerodromiinae, Clinocerinae, Tachydromii- terms of generic diversity, the entire group is
nae and Hybotinae) while others gather in aerial richest in western North America, southern
mating swarms (Atelestinae, Ocydromiinae, Africa, New Zealand and southern South
most Empidinae, and many Trichopezinae). The America. This is also reflected in species diver-
synchronized movement of adult flies within sity, where non-dolichopodid Empidoidea are
these mating swarms is the basis for the com- seemingly more diverse in temperate rather than
mon name “dance flies.” In a few genera, such as tropical regions.
Trichina (Trichininae), Oedalea (Oedaleinae) The Empididae are a large family of well over
and Bicellaria (Hybotinae), aerial aggregations 2,000 described species worldwide that include
have been observed even though mating appar- the Empidinae, Hemerodromiinae, and Clinoceri-
ently takes place on the ground. These aerial nae, as well as the Ragas group, Hesperempis group,
aggregations that occur without mating have Philetus and Brochella. The monophyly of this
been termed “relict swarms.” diverse heterogeneous family is still somewhat
Members of one large subfamily, the uncertain. The immature stages of Clinocerinae
Empidinae, transfer nuptial gifts from male to and most Hemerodromiinae are aquatic whereas
female during courtship and mating. Depending the immature stages of the other groups are pri-
on the species, these nuptial gifts include prey, marily terrestrial.
various types of inedible objects, and/or secreted The Atelestidae are a small distinctive family
balloons (Fig. 1a), which is why the entire group of nine extant species that are classified in four
is sometimes referred to as “balloon flies.” Within genera. The Palaearctic genus Nemedina is assigned
the Empidinae mate choice is generally made by to its own subfamily Nemedininae whereas the
females that visit male-dominated swarms. Males other genera Atelestus (Palaearctic), Meghyperus
in these species compete for females using these (Holarctic) and Acarteroptera (Chilean) belong
nuptial gifts, often enhanced with various to the Atelestinae. The immature stages of
secondary sexual characters. However, many Atelestidae are unknown, but are believed to be
species of Empidinae exhibit sex-role reversed terrestrial.
courtship behavior where females gather in The Hybotidae are a diverse family of approxi-
swarms to await males that choose their mates. mately 2,000 described species currently classified
Females of these species exhibit many different into five subfamilies, namely the Trichininae,
secondary sexual characters used in courting Ocydromiinae, Oedaleinae, Tachydromiinae, and
males, such as enlarged wings, pinnate leg scales, Hybotinae. The family is well defined by several
eversible abdominal pleural sacs and silvery synapomorphies and its monophyly is not in doubt.
abdomens. All of the included subfamilies are virtually
D
Darkling Beetles (Coleoptera: Tenebrionidae) 1151
Dark Mealworm, Tenebrio
cosmopolitan in distribution, except for Trichini-
obscurus Fabricius (Coleoptera:
nae and Oedaleinae, which are essentially Holarc-
Tenebrionidae)
tic. The entire family is terrestrial and many species
are found in some of the driest habitats occupied
by empidoid flies. This is a stored grain insect, and very similar to the
The Brachystomatidae are a moderate-sized more common yellow mealworm, Tenebrio moli-
family of about 150 described species worldwide. tor Linnaeus.
The family includes the nearly cosmopolitan  Stored Grain and Flour Insects
Trichopezinae with 14 recognized genera, the  Darkling Beetles
Southern Hemisphere Ceratomerinae with three
currently recognized genera, and the primarily
Darkling Beetles (Coleoptera:
Holarctic Brachystomatinae with three genera.
Tenebrionidae)
The monophyly of this morphologically diverse
family is based primarily on synapomorphies
associated with modifications of the female ter- charles a. triplehorn
minalia. Based on limited information obtained The Ohio State University, Columbus, OH, USA
from rearing records and habitats occupied by
adults, the immature stages of Brachystomatidae The family Tenebrionidae belongs to the suborder
are believed to include both terrestrial and aquatic Polyphaga of the order Coleoptera (beetles). The
representatives. superfamily Tenebrionoidea includes the families
 Flies Anthicidae, Colydiidae, Melandryidae, Monomi-
dae, Salpingidae, Mycetophagidae, Tetratomidae,
Mordellidae, Rhipiphoridae, Prostomidae, Syn-
References chroidae, Oedemeridae, Stenotrechelidae,
Meloidae, Mycteridae, Ciidae, Boridae, Pythidae,
Aderidae, Scraptiidae, Archeocrypticidae, and
Chvála M (1983) The Empidoidea (Diptera) of Fennoscandia
and Denmark. II. General Part. The families Hybotidae, Zopheridae, as well as the large family
Atelestidae and Microphoridae. Fauna Entomol Scand Tenebrionidae.
12:1–279
Order: Coleoptera
Collin JE (1961) Empididae. British flies, vol 6. University
Press Cambridge, UK, 782 pp Suborder: Polyphaga
Cumming JM (1994) Sexual selection and the evolution of Superfamily: Tenebrionoidea
dance fly mating systems (Diptera: Empididae; Empidi-
Family: Tenebrionidae
nae). Can Entomol 126:907–920
Tenebrionidae are the fifth largest family of
Cumming JM, Sinclair BJ (2007) Family Empididae. In: Brown
BV, Borkent A, Cumming JM, Wood DM, Woodley Coleoptera with 14,641 species worldwide and
NE, Zumbado M (eds) Manual of Central American 1,345 occurring in the United States. These num-
Diptera, vol 1. NRC Research Press, Ottawa, Canada (in
bers are low since in the past three decades, many
press)
changes have been made in the classification of
Sinclair BJ, Cumming JM (2006) The morphology, higher-
level phylogeny and classification of the Empidoidea the family and many new taxa have been described.
(Diptera). Zootaxa 1180:1–172 The Zopheridae and Archeocrypticidae have been
Smith KGV (1969) The Empididae of southern Africa (Dip-
removed and given family status, and the former
tera). Annals Natal Mus 19:1–342
Steyskal GC, Knutson LV (1981) Chapter 47. Empididae. In: families Alleculidae and Lagriidae are now con-
McAlpine JF, Peterson BV, Shewell GE, Teskey HJ, sidered subfamilies within Tenebrionidae. Adding
Vockeroth JR, Wood DM (eds) Manual of Nearctic
species from those two groups raises the total
Diptera, vol 1. Agriculture Canada Monograph 27,
world species to 16,267 with 1,540 in the United
Biosystematics Research Institute, Ottawa, Canada,
674 pp States. The current classification recognizes twelve
D
1152 Darkling Beetles (Coleoptera: Tenebrionidae)
subfamilies worldwide, eleven of which occur in their resemblance to the larvae of Elateridae which
the United States. are known as “wireworms.” Both do similar damage
Tenebrionidae is one of the most highly to underground portions of plants. The most con-
evolved and diverse families of Tenebrionoidea spicuous difference between them is that tenebri-
and, indeed, in the order Coleoptera itself. It is a onid larvae have a less flattened head and unfused
difficult group to characterize because of the labrum. Larvae of elaterids usually have the labrum
many exceptions that exist in most characters. absent or fused with the clypeus, forming a median
Discounting the exceptions, the following com- sclerotized projection called a “nasale.”
bination of morphological characters separate
adult tenebrionids from other Tenebrionoidea:
Habitats
tarsal formula 5-5-4; procoxal cavities closed;
antennae eleven-segmented and inserted under
lateral expansion of the genae; abdomen with There are few macrohabits from which tenebrion-
basal three visible sterna connate, 4 and 5 ids are excluded. In the deserts of the world, they
movable. are among the most conspicuous occupants, some-
times occurring in enormous numbers. All of
them have evolved remarkable water conserving
Adults
mechanisms, and survive principally on metabolic
water derived from their foods. Essentially all of
The length of adult tenebrionids ranges from these desert dwellers lack metathoracic wings and
slightly more than one to more than 60 mm. While are therefore flightless. The elytra usually are fused,
the popular concept of the family is somber color- forming a subelytral cavity, and the integument is
ation (shades of black or brown), there are some thick, both factors helping to control water loss
(i.e., Strongylium, Cuphotes, Poecilisthus, Pla- through the spiracles.
tydema) that are vividly colored. There are many Desert tenebrionids escape the extreme day-
species in a number of different subfamilies which time temperatures of arid regions by hiding under
lack metathoracic wings (sometimes there are debris or in mammal burrows, venturing out only
apterous and alate members within the same at dusk. They are probably not truly nocturnal but
genus, and occasionally even within a population opportunists, taking advantage of cooler night-
of a single species (i.e., Blapstinus). The elytra usu- time temperatures. The experienced darkling bee-
ally completely cover the abdomen but in some tle collector works mostly at night.
(i.e., Corticeus, Phaleria) the pygidium is exposed. At the other extreme are many genera, rich
in species, which inhabit moist environments
throughout the world. Genera such as Strongylium,
Immature Stages
Helops, Tarpela and Statira are almost entirely
confined to tropical and subtropical regions and
Tenebrionid larvae are usually elateroid (wireworm- are more diurnally active.
like) or C-shaped, elongate and cylindrical, 5–40 Several genera in the tribe Phaleriinae (e.g.,
mm in length, hard-bodied, with five-s egmented Phaleria, Chaerodes) are found only on ocean
thoracic legs. The head is exserted with a frontocly- beaches where they congregate beneath organic
peal suture present. The apical abdominal segment debris. There are a number of interesting rela-
is usually rounded or triangular, urogomphi, con- tionships between tenebrionids and other insects.
cave plates, or other armature present or absent. Members of the genus Corticeus inhabit the gal-
The larvae of some tenebrionids (e.g., Eleodes) leries of bark beetles (Scolytinae) and others (e.g.,
are sometimes called “false wireworms” because of Araeoschizus, Bycrea) occur in ant nests.
D
Darlington, Jr., Philip J 1153
Many species are subcortical but usually Doyen JT, Lawrence JF (1979) Relationships and higher clas-
sification of some Tenebrionidae and Zopheridae
occur under bark of a number of tree species.
(Coleoptera). Syst Entomol 4:333–377
There are few that are host specific and none are
Doyen JT, Tschinkel WR (1982) Phenetic and cladistic rela-
defoliators. At least one species, Palembus ocularis tionships among tenebrionid beetles (Coleoptera). Syst
Casey, appears to be intimately associated with a Entomol 7:127–183
Watt JC (1974) A revised subfamily classification of Tenebri-
plant, Tamarindus indica, and another, Sciophagus
onidae (Coleoptera). NZ J Zool 1:381–452
pandanicola (Boisduval), occurs on Pandanus in
Watt JC (1967) A review of classifications of Tenebri-
,
Hawaii. Those are exceptions. onidae (Coleoptera). Entomologists Monthly Maga-
There are a number of genera in several widely zine 102:80–86
separated phylogenetic groups (e.g., Bolitotherus,
Rhipidandrus, Neomida, Platydema) which inhabit
Dark-Winged Fungus Gnats
fungi. It appears that both the larvae and adults
feed directly on the fungi.
A number of darkling beetle species live in Members of the family Sciaridae (order Diptera).
caves or rock fissures and apparently nowhere  Flies
else. Some (e.g., Alphitobius laevigatus (Fab.),
Zophobas spp.) often are associated with bat
Darlington, Jr., Philip J
guano in caves.
Philip Darlington was born in Philadelphia, USA,
Food
on November 14, 1904. The family moved to the
state of Connecticut, and Philip spent the rest of his
Darkling beetles are mostly omnivorous and only life in the New England states (Connecticut, Massa-
a few appear to be host specific. Many species (e.g., chusetts, Maine, New Hampshire and Vermont)
Eleodes, Blapstinus) are root feeders. Wherever except when he was on field trips or in military ser-
stored grain products are stored, a number of cos- vice. His parents were interested in nature, which
mopolitan tenebrionids are secondary pests. None influenced Philip in his career. In 1922, he entered
of them are able to attack unbroken grain kernels Harvard University, gained a first degree and contin-
but feed on ground or milled farinaceous materi- ued to graduate school, gaining a Ph.D. in 1932. An
als; these insects are collectively known as “bran explanation for his lengthy studies is that he took the
bugs.” The genera Tribolium, Gnatocerus, Palorus, year 1928–1929 to work in Colombia for the United
Sitophilus and Latheticus are secondary pests of Fruit Company. Ostensibly he worked as entomolo-
cereal products throughout the world. All of these gist to solve a problem in applied entomology. He
are thought to be of tropical origin and probably returned to Harvard University with a large collec-
were fungus feeders originally. They tend to favor tion of insects and bird skins. Then he worked on
stored products that are not in the best condition his Ph.D. dissertation on carabid beetles of New
(wet, moldy, etc.). Hampshire. But in 1931–1932 he joined an expedi-
tion, led by W.M. Wheeler, to Australia, from which
he returned with an extraordinarily large collection
of insects plus 341 specimens of mammals. On his
References
return to Harvard University, he was appointed
assistant curator of insects. During the next few
Doyen JT (1972) Familial and subfamilial classification of the
years he made several expeditions to the West Indies,
Tenebrionoidea (Coleoptera) and a revised generic clas-
sification of the Coniontini (Tentyriinae). Quaestiones collecting insects and other animals in the high
Entomol 8:357–576 mountains of the Greater Antilles. In 1940 he was
D
1154 Darners
Darwin, Charles
appointed curator of Coleoptera, and 2 years later he
married Elizabeth Koch. However, in World War II
he enlisted in the U.S. Army and served in a malaria Charles Darwin was born in Shrewsbury, England,
survey in New Guinea, the Bismarck Archipelago, on February 12, 1809. He was educated at Shrews-
and the Philippines. He returned to Harvard Uni- bury Grammar School, Edinburgh University
versity in 1944. Other collecting expeditions fol- (1825–1827), and Cambridge University
lowed, such as to the southern extremes of South (1828–1831), obtaining B.A. and M.A. degrees. His
America in 1962–1963, but he was also appointed real interest at Cambridge was in collecting insects,
Alexander Agassiz Professor of Zoology at Harvard especially beetles. Then he served as naturalist
University. His published works were on zoogeogra- aboard H.M.S. Beagle during that ship’s round-t he-
phy and natural selection, mimicry, and taxonomy. world cruise in 1831–1836. Insects were prominent
His three books were (1957) “The geographical dis- among the organisms that he observed and c ollected,
tribution of animals,” (1965) “Biogeography of the but many of the new species that he collected were
southern end of the world…,” and (1980) “Evolution not named by him, but by specialists (C.C. Babing-
for naturalists…” In recognition of his works he was ton, F.W. Hope, W.W. Saunders, J.O. Westwood, and
elected to the (U.S.) National Academy of Sciences, G.R. Waterhouse). He did spend 8 years
to the American Academy of Arts and Sciences, and (1846–1854) monographing barnacles (Cirripedia).
was awarded two Guggenheim fellowships. He died His leaving of the descriptive work on insects to
on December 16, 1983. specialists allowed him to concentrate on produc-
ing his greatest works, “The origin of species” (1859)
and “The descent of man” (1871). He died on April
References
19, 1882, in the county of Kent.
Ball GE (1985) The contributions of Philip J. Darlington, Jr. to
the study of carabid beetles in the Americas and a References
bibliography of his publications. In: Ball GE (ed)
Taxonomy, phylogeny and zoogeography of beetles and
ants. A volume dedicated to the memory of Philip Remington JE, Remington CL (1961) Darwin’s contributions
Jackson Darlington, Jr. (1904–1983). Dr W. Junk (Series to entomology. Ann Rev Entomol 6:1–12
Entomologica vol 33), Dordrecht, xiii + 514 pp Smith KGV (1982) Charles Darwin and the Royal Entomo-
Brown WL (1985) Philip Darlington’s contributions to evolu- logical Society of London. Antenna 6:200–201
tionary theory. In: Ball GE (ed) Taxonomy, phylogeny
and zoogeography of beetles and ants. A volume dedi-
cated to the memory of Philip Jackson Darlington, Jr.
(1904–1983). Dr W. Junk (Series Entomologica vol 33), Darwinism
Dordrecht, xiii + 514 pp
Carpenter FM (1985) Philip Jackson Darlington, Jr.: A bio-
graphical sketch. In: Ball GE (ed) Taxonomy, phylogeny This term is synonymous with the Theory of Natu-
and zoogeography of beetles and ants. A volume dedi- ral Selection, which explains that speciation results
cated to the memory of Philip Jackson Darlington, Jr.
from differential survival and reproduction in
(1904–1983). Dr W. Junk (Series Entomologica vol 33),
response to selective forces (both biotic and abi-
Dordrecht, xiii + 514 pp
otic). Organisms that are well adapted to selective
forces can survive and reproduce, with the prog-
Darners
eny of these survivors likely inheriting these adap-
tations. In contrast, organisms that are not well
A family of dragonflies in the order Odonata: adapted to these same selective forces may not
Aeshnidae. survive or reproduce, leading to their elimination
 Dragonflies and Damselflies and loss of progeny. Thus, small mutations interact