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 Fig. 1. Scheme showing visible stages of development in a typical endopterygote insect, emphasizing immature stages.
(e.g. Danks, 1971 for chironomids; Wiggins, 1973 for
limnephilid caddisflies).
Larval stage. The major role of larvae in the life cycle
is to feed and grow. Consequently, their structural and
temporal patterns of development are related largely to
the need to survive in particular environments (generally
chosen by ovipositing females) and to acquire and use the
food resources there.
In many species first instars differ from later instars,
especially if they serve for dispersal (as in the planktonic
first instars of Chironomidae: Davies, 1976), host finding
(as in parasitoids that penetrate hosts from the host
habitat) or protection of the resource from other indi-
viduals (as in the hyper-metamorphic first instars of many
parasitic hymenopterans). Typically, first instars moult
relatively rapidly to the second instar, and in some species
do not feed [Harvey, 1957 for the tortricid moth Choristo-
neura fumiferana (Clem.); Strickman et al., 1997 for the
pholcid spider Crossopriza lyoni (Blackwall)]. Depending
on the group or species and on its particular life cycle,
other instars are modified. For example, the pre-adult
(second nymphal) instar in several Acaridiae is the
hypopus, a modified resting instar.
The number of instars typically is large in apterygotes
(e.g. up to 30 in silverfish: Nishizuka et al., 1998) and
primitive exopterygotes. Mayflies normally have 1525
instars, but from as few as 10 to as many as 52 depending
on the species (Trost & Berner, 1963; Clifford et al.,
1979; Fink, 1980; Brittain, 1982). There are about 1224
instars in stoneflies (Vaught & Stewart, 1974; Baumann,
1987), and 1015 in dragonflies (Westfall, 1987). On the
other hand, usually there are only five instars in Heterop-
tera. Advanced endopterygotes typically have only 34
instars (many Coleoptera and most Diptera) or 47 instars
(most Lepidoptera and Hymenoptera). Although in many
species, especially of endopterygotes, the number of
instars is fixed, in some species the number can vary
depending on the conditions of temperature and food
experienced. For example, variations in the number of
instars are known in spiders (Miyashita, 1997), ticks
(Dautel & Knlle, 1997), mayflies (Brittain, 1976), stone-
286
flies (Vaught & Stewart, 1974), dragonflies (Rivard &
Pilon, 1977), grasshoppers (Kawano & Ando, 1997), bee-
tles (Beck, 1971; Gerard & Ruf, 1997), moths (Neunzig,
1969; Morita & Tojo, 1985; Clare & Singh, 1990;
Kamata & Igarashi, 1995), and other groups. The devel-
opmental routes of individuals with different numbers of
instars are not equivalent, and data on instar durations
have to be recorded separately for these different indi-
viduals.
At any given time, each instar can be in one of three
states: active and feeding; inactive and moulting; or inac-
tive and dormant. Moults can occupy a significant frac-
tion of the time spent in the larval period, especially at
lower temperatures; e.g. 43% of the penultimate instar in
the geometrid moth Epirrita autumnata (Borkhausen) and
50% in the chrysomelid beetle Galerucella sagittariae
Gyllenhal at 12C (Ayres & Maclean, 1987). Moreover,
larval moults, pupation and adult eclosion are prevented
at low temperatures that are otherwise suitable for growth
and development.
Dormancies and related responses can greatly prolong
life-cycle durations. Many life-cycle programmes are
structured by diapauses that suppress larval development
in response to environmental cues of various kinds
(Danks, 1987). Beyond such clearly marked develop-
mental delays, the rate of growth can be modified indi-
rectly by photoperiod (Danks, 1987, Table 3); and simple
quiescence can be caused directly by unsuitable food or
by conditions that are too cold or too dry (see Delays).
The possibility of such interruptions in the life cycle
prompts care in developing experimental protocols to
measure life-cycle duration.
Once feeding ends, the final-instar larva enters the pre-
pupal stage (Fig. 1). A few authors have restricted the use
of this term, like the term propupa for Thysanoptera, to a
separate preadult instar. The end of feeding can some-
times be detected by direct observation, although this is
often inadequate for rapid checking during experiments.
It is also signalled by changes in behaviour or by changes
in appearance. For example, fully grown larvae may spin
a cocoon, or leave the food to wander in search of a pupa-
 Fig. 2. Scheme showing visible stages of development in a typical oviparous insect, emphasizing adult stages.
tion site (Richard et al., 1986). Prepupae normally have
thickened or more conspicuous anterior segments, corre-
sponding to the thorax of the pupa, which are easily
visible in the larvae of Diptera such as Chironomidae, for
example. Duration of the prepupal stage, unlike that of
the larva that precedes it, does not depend directly on
food, but instead may be determined partly by the avail-
ability of suitable pupation sites as well as by
temperature.
Pupal stage. Substages of pupal development can be
recognized from selected characters such as leg or wing
rudiments, eyespots or other pigmentation, thoracic or
abdominal setae, and internal reproductive structures.
Colour darkens markedly as eclosion nears. The charac-
teristic U-shaped respiration curve of pupae shows
decreased metabolism after pupation and a larger increase
before eclosion (Schneidermann & Williams, 1953).
The substages of pupal development have not been
characterized in most species, although some diapause
stages or markers have been documented (e.g. Ohnesorge,
1979; Hackett & Gatehouse, 1982; Kusters & Herrebout,
1989).
Adult stage. Several relatively easily recognized sub-
stages of adult life are important for assembling data on
the life cycle (Fig. 2). These substages are generally
visible in mites as well as in insects. Only Collembola
and other primitive hexapods continue to moult during
adult life.
Upon eclosion from the last-instar larva or pupa, the
cuticle is still soft, and the adult is teneral. Adults harden
very rapidly in some species, but in others the process of
sclerotization may take many days, as in dragonflies. In
species that chew their way out of a pupal cocoon, mandi-
bles must harden before this is possible, and therefore
emergence is delayed for some days after adult eclosion
(see Hidden stages).
Information on oviposition is very important for under-
standing the duration of the life cycle: the life cycle of a
typical individual begins when it is deposited as an egg,
but determining the corresponding endpoint depends on
the temporal pattern in which eggs of the next generation
are produced. In most species mating alters preoviposi-
tion or oviposition periods or longevity (see Laboratory
rearing).
In many species the preoviposition period is longer than
the teneral period, the premating period or both, for two
reasons. First, substantial ovarian development, often pre-
ceded or accompanied by adult feeding, may be required.
For such species, the supply of adult food helps to deter-
mine the pattern of oviposition (see also Laboratory rear-
ing). Second, many species include a period of dispersal
early in the life of some or all adults before eggs are
developed (see Johnson, 1969; Dingle, 1972 for general
information on this oogenesis-flight syndrome). In
polymorphic species (Zera & Denno, 1997), the develop-
mental pathway usually differs between flying and non-
flying individuals. Long-distance flight takes place before
and after reproductive diapause in some species (e.g.
Oku, 1983), and a similar phenomenon on a smaller scale
is common in many other insects (Wissinger, 1997). In
such species it may be difficult to assess oviposition
responses under laboratory conditions.
To summarize oviposition, its temporal pattern as well
as the beginning and end must be described. Normally,
more eggs are laid soon after oviposition begins than at
the end of the oviposition period. A few species oviposit
in two or more separate bouts, such as mosquitoes that
mature egg batches after successive meals, and long-lived
adults that survive for more than one season (Danks,
1992).
Some other species, especially those with long-lived
adults, survive for some time before they die without
depositing any more eggs (Fig. 2). Such postoviposition
adults have been noted in many laboratory studies, and
spent females have also been reported under field con-
ditions (e.g. Matalin, 1998). The fact that some adults sur-
vive after oviposition ends, and the fact that the
distribution of oviposition through the oviposition period
is usually skewed, means that the commonly measured
287
intervals of preoviposition period and longevity are
inadequate on their own to characterize oviposition.
Omitted or abbreviated stages. Various groups or
species have reduced or eliminated some of the character-
istic life-cycle stages shown in Figs 1 and 2. Such
changes may serve to remove or protect vulnerable stages
or to accelerate the life cycle. Apart from reduction in the
number of instars, one of the commonest changes is
reduction of eggs through larviposition or viviparity, as in
tachinids, some sarcophagids and aphids. Eggs or larvae
or both can be retained in the mother, even resulting in an
adult to adult life cycle, as in ovoviviparous pyemotoid
mites (Wrensch & Bruce, 1991). A few parthenogenetic
species can produce offspring even before adult eclosion,
as in the aphid Schizaphis graminum (Rondani) (Wan-
jama & Holliday, 1987), and in some individuals of the
black fly Prosimulium ursinum (Edwards) (Carlsson,
1962 as Simulium). Non-feeding instars also occur com-
monly (e.g. some first instars as already noted; special-
ized dormant stages). Adult stages are abbreviated by
features such as parthenogenesis (mating is unnecessary),
adult eclosion with eggs fully developed (ovarian devel-
opment is not necessary), and rapid oviposition in a single
batch.
The many differences of this sort make it more difficult
to compare the durations of different stages and substages
among species. Care must be taken not to compare or
average statistics for substages that are not equivalent.
Hidden stages. In practice, not all of the moults
between successive stages are readily observed, so that
statistics on duration are much more difficult to collect.
For example, egg hatch cannot be observed directly in
gall formers, endoparasitoids and other species that live
inside their food resources. The prepupal to pupal change
is not visible without special procedures in species that
pupate inside robust silken cocoons, or in dense sub-
strates such as soil. In cyclorrhaphan Diptera, pupation
takes place inside the puparium (the rounded off, sclero-
tized exoskeleton of the third-instar larva) up to a few
days after pupariation, and hence normally can be deter-
mined only by dissection.
Fig. 3. Stylized cumulative event curve and event line, illus-
trating means to summarize key measures during the interval
over which an event occurs in the population. For further expla-
nation, see text.
288
The pupal to adult change is difficult to observe
directly in species that remain quiescent in a pupal cell or
relatively robust cocoon after eclosion and while still
teneral. Adult escape or emergence requires up to
several days after eclosion in species of chrysomelids,
curculionids, and other beetles (e.g. Buckingham & Ben-
nett, 1981; Jackson & Elliott, 1988; Purcell & Balcuinas,
1994) and in various Hymenoptera (e.g. Danks, 1970, p.
332). Additional delays occur if the adults subsequently
are dormant, as in xylocopine bees of the genus Ceratina
(e.g. Maeta et al., 1992). The post-eclosion interval can
be estimated in adults isolated from their cocoons by the
fact that meconium normally is discharged when exit
from the pupal cell would have occurred.
Sexual differences. In many species rates of develop-
ment differ between the sexes. Conventional wisdom sug-
gests that males develop more rapidly because they are
smaller or because protandry may be advantageous for
mating, whereas females develop more slowly because
they are larger or can increase fecundity by further
feeding (cf. Danks, 1994). However, this pattern is by no
means the only one (Honk, 1997).
MEASURES OF DURATION
Temporal programmes of development or life-cycle
pathways of many species are relatively complex, so that
no simple set of universal experiments or measurements
can give unambiguous information about substage dura-
tions in every species. This section briefly provides some
context for the choice and recording of intervals, and also
considers developmental delays, which complicate assess-
ments of duration in many species.
Temporal patterns and key measures. Two types of
temporal information can be recorded about the life cycle
of an individual. First, each individual experiences unique
events, such as hatch and pupation, which together
delimit larval duration. Second, patterned events may take
place within a particular interval, notably the pattern of
oviposition within an individual oviposition period
(Danks, 1994), which is averaged to delimit the end of
the current generation and the start of the next. Such pat-
terns can be described in part by means of summary sta-
tistics, but accurate documentation of the pattern is com-
plex.
Summary data such as the date by which 50% of the
eggs are deposited can be obtained either by averaging
for the population as a whole (the date on which 50% of
the eggs have been deposited by the whole population) or
by averaging individual records (the mean of the dates on
which each individual has deposited 50% of its eggs).
These values are not always the same; the former is nor-
mally preferable.
Key statistics include the first, 50%, median, and last
occurrences. Especially when the temporal pattern is
more or less normally distributed (and hence sigmoid in a
cumulative plot), valuable information is also provided by
the 10% and 90% levels, which correspond closely to the
inflections of the sigmoid curve (see Fig. 3), and it is fea-
sible to express such information using marked lines (top
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