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Field
Guide to Common Western Grasshoppers
by Robert
E. Pfadt
Introduction
| External Anatomy |
Names and Species | Grasshopper Populations
| Life Cycles | Seasonal
Occurrence and Behavior | Collections
and Survey | Grasslands of North America
| Food Plants | Glossary
| Selected References |
Species Fact Sheets | (PDF)
Seasonal
Cycles
An important component of grasshopper life history is the seasonal
cycle - the timing of the periods of egg hatch, nymphal growth and
development, emergence of the adults and acquisition of functional
wings (fledging), and the deposition of eggs or reproduction. The
occurrence of these periods varies among the species and is greatly
influenced by weather. An early spring hastens these events and
a late one delays them. Latitude also influences the dates of occurrence.
In North America springtime comes earlier in the south and later
in the north. Consequently, hatching, development and maturation
come earlier in the south and later in the north. In the West, altitude
is also an important factor. The lower temperatures of higher altitudes,
especially those of mountain meadows, are responsible for retarded
seasonal cycles of grasshoppers and may often cause a two-year life
cycle among species that ordinarily have a one-year life cycle.
Nymphal and adult grasshoppers are present all year long in natural
habitats. Several species overwinter in late nymphal stadia and
become adult in early spring. The majority, however, pass the winter
as eggs protected in the soil. Depending on the species, these eggs
hatch at different times from early spring until late summer. The
variety of seasonal cycles allows actively feeding and developing
grasshoppers to spread out over the entire growing season. Each
species has its own time to hatch, develop, and reproduce, but with
much overlapping of the cycles. An experienced scout going into
the field to survey expects to find certain species at certain times
and is thus aided in making identifications. Table 5 arranges the
seasonal cycles of grasshoppers into: (1) very early nymphal and
adult group, (2) very early hatching group, (3) early hatching group,
(4) intermediate hatching group, and (5) late hatching group. The
table is especially helpful in the identification of young nymphs.
| TABLE 5.
Species of grasshoppers grouped by seasonal appearances. |
|
SLANTFACED
(Gomphocerinae)
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BANDWINGED
(Oedipodinae)
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SPURTHROATED
(Melanoplinae)
|
|
Early
spring, large nymphs and adults. Overwintered in nymphal stage.
|
|
Eritettix simplex
Psoloessa delicatula
Psoloessa texana
|
Arphia conspersa
Chortophaga viridifasciata
Pardalophora apiculata
Pardalophora haldemani
Xanthippus corallipes
Xanthippus montanus
|
. |
|
Very
early hatching group, hatching in early spring.
|
|
Aeropedellus clavatus
|
. |
Melanoplus confusus
|
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Early
hatching group, hatching in mid-spring.
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|
Acrolophitus hirtipes
Ageneotettix deorum
Amphitornus coloradus
Aulocara elliotti
Cordillacris occipitalis
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Camnula pellucida
Dissosteira pictipennis
Dissosteira spurcata
Trimerotropis pallidipennis
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Aeoloplides turnbulli
Melanoplus angustipennis
Melanoplus bivittatus
Melanoplus cuneatus
Melanoplus foedus
Melanoplus infantilis
Melanoplus occidentalis
Melanoplus packardii
Melanoplus rugglesi
Melanoplus sanguinipes
Oedaleonotus enigma
|
|
Intermediate
hatching group, hatching in late spring.
|
|
Aulocara femoratum
Boopedon nubilum
Chorthippus curtipennis
Cordillacris crenulata
Mermiria bivittata
Orphulella speciosa
Parapomala wyomingensis
Phlibostroma quadrimaculatum
Pseudopomala brachyptera
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Derotmema haydeni
Dissosteira carolina
Dissosteira longipennis
Hadrotettix trifasciatus
Mestobregma plattei
Metator pardalinus
Spharagemon collare
Spharagemon equale
Trachyrhachys kiowa
Trimerotropis campestris
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Hesperotettix viridis
Hypochlora alba
Melanoplus bowditchi
Melanoplus dawsoni
Melanoplus devastator
Melanoplus differentialis
Melanoplus flavidus
Melanoplus femurrubrum
Melanoplus marginatus
Brachystola magna
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Late
hatching group, hatching begins in early summer.
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Eritettix simplex
Opeia obscura
Psoloessa delicatula
Psoloessa texana
Stenobothrus brunneus
Syrbula admirabilis
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Arphia conspersa
Arphia pseudonietana
Chortophaga viridifasciata
Pardalophora apiculata
Pardalophora haldemani
Xanthippus corallipes
Xanthippus montanus
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Dactylotum bicolor
Melanoplus alpinus
Melanoplus borealis
Melanoplus bruneri
Melanoplus keeleri
Melanoplus lakinus
Phoetaliotes nebrascensis
Schistocerca emarginata
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Behavior
A grasshopper's day (and night) are linked closely with the physical
factors of the environment, especially temperature, but also light,
rain, wind, and soil. Stereotyped and instinctive behavior patterns
serve grasshoppers remarkably well in making adjustments to wide
fluctuations of physical factors that otherwise might be fatal.
Grasshoppers effectively exploit the resources of their habitat
and at the same time are able to tolerate or evade the extremes
of physical factors. Their characteristic rapid jumping and flying
responses help them escape numerous enemies that parasitize or feed
upon them.
In temperate North America, certain behavior patterns are held
in common among grasshopper species, especially among those occupying
the same part of the habitat, while other patterns differ. Individuals
of different species have different ways of spending the cool nights.
Some retreat under litter or canopies of grasses; others squat on
bare ground and take no special shelter. Still others may climb
a small shrub or a tall grass plant and rest at various heights
within the canopy. Under favorable conditions of temperature and
other elements of weather, grasshoppers may be active and even feed
during the night. In southwestern states they have been observed
on warm nights wandering about on the ground and on vegetation,
feeding, and stridulating. Several species have been recorded flying
at night and are attracted to city lights. A temperature of 80°F
is apparently a prerequisite for night flying with maximum flight
activity occurring at temperatures above 90°F.
A grasshopper's day usually starts shortly after dawn. Because
body temperatures have fallen during the night, a grasshopper on
the ground crawls to an open spot, often on the east side of vegetation,
that allows it to warm itself by basking in the radiant rays of
the sun. A common orientation is to turn a side perpendicular to
the rays and lower the associated hindleg, which exposes the abdomen.
Those that have spent the night on a plant make adjustments in their
positions to take advantage of the sun's rays or they may climb
or jump down to the ground to bask. Although grasshoppers generally
remain quiet while they bask, they occasionally stir, preen, turn
around to expose the opposite side, and sometimes crawl to a more
favorable basking location. Grasshoppers may bask for a second time
in the cool of late afternoon. Then as shadows begin to engulf the
habitat, they retreat into their customary shelters.
After basking for one to two hours on sunny days, grasshoppers
become active. They may walk about, seek mates, or feed. Because
grasshoppers are cold-blooded creatures, their usual daily activities
are interrupted when the weather turns cold, overcast, or rainy.
During such times they generally remain sheltered and inactive.
During warm sunny weather of late spring and summer, grasshoppers
take advantage of two foraging periods, one in the morning and one
in the afternoon. Different species of grasshoppers have different
ways of attacking and feeding on their host plants. Individuals
of certain species climb the host plant to feed on leaves, petals,
buds, or soft seeds while others cut and fell a grass leaf and feed
on it while sitting on the ground. Geophilous (ground-dwelling)
species regularly search the ground for food, picking up and feeding
on seeds, dead arthropods, and leaves felled by other grasshoppers.
In grasslands with considerable bare ground, very little litter
is produced by grasshoppers. Severed leaves, dropped by the grasshoppers
that feed on the host plant, are soon found and devoured by the
ground foragers. In habitats infested by dense populations of grasshoppers,
pellets of their excrement, rather than litter, accumulate in small
conspicuous piles. Only in habitats of tall grass do grasshoppers
produce leafy ground litter that goes uneaten and "wasted."
This is because the grasshopper species in these habitats are phytophilous
and feed resting on the host plant.
Grasshoppers fastidiously select their food. By lowering their
antennae to the leaf surface and drumming (tapping) it with their
maxillary and labial palps, grasshoppers taste a potential food
plant. Gustatory sensilla located on the tips of these organs are
stimulated by attractant and repellent properties of plant chemicals,
allowing a grasshopper to choose a favorable host plant and reject
an unfavorable one. A grasshopper may take an additional taste by
biting into the leaf before it begins to feed freely. Phagostimulants
are usually important ones nutritionally-certain sugars, phospholipids,
amino acids, and vitamins. Grasshoppers may even make choices among
the leaves of a single host plant. They prefer young green leaves
and discriminate against old yellowing ones. Nevertheless, individuals
of ground-dwelling species often feed in short bouts on old plant
litter lying on the ground as well as on dry animal dung. This feeding
may be a means of restoring water balance - either losing or gaining
moisture.
Later, in the species fact sheets, grasshopper food plants are
referred to by their common names. The appendix (Grasshopper
Food Plants) provides a listing of these with their scientific
names to clarify any ambiguities.
Amazingly, grasshoppers are able to communicate visually and acoustically
among themselves. They produce sounds with structural adaptations
on hindlegs and wings and receive these signals with auditory organs
(ears) located in the first abdominal segment (Fig. 8). Using their
colorful wings and hindlegs they also flash visual messages and
receive these with their compound eyes. Intraspecific communication,
that which occurs between members of the same species, is used to
attract and recognize mates, to ward off an unwanted suitor, and
to defend a territory or a morsel of food. Grasshoppers produce
acoustical signals by rubbing the hindlegs against the tegmina (Fig.
12) or the sides of the abdomen. They may also communicate by rapidly
flexing or snapping their hindwings in flight, a behavior called
crepitation. Each species apparently produces its own unique
sound and, in human terms, has its own language. The details, where
known, of finding a mate are described in the species section of
this guide.

Figure 12. Signals and
postures commonly occurring in grasshoppers: (a) ordinary stridulation
-- a slow, stereotyped, repetitive or non-repetitive, high amplitude
movement in which the femur rubs against the forewing; (b) vibratory
stridulation -- a fast, stereotyped, repetitive, low amplitude
movement in which the femur rubs against the forewing; (c) ticking
-- a stereotyped, repetitive or non-repetitive movement in which
the tibiae are kicked out and struck against the ends of the forewings;
(d) femur-tipping -- a silent, stereotyped, non-repetitve
raising and lowering of the femora; (e) Femur-shaking (with
substrate-striking) -- a stereotyped, repetitive shaking of the
femora in which ends of the tibiae strike the substrate to produce
substrate vibration or a drumming sound; (f) femur-shaking
(silent or with wing-striking) -- similar to (e) but the movement
is silent, or the femora strike the forewings; (g) femur-raising
-- a slow, graded non-repetitive movement that may or may not be
accompanied by mild upward kicking motions of the tibiae; (h) presenting
-- a variable, graded posturing of the female in response to male
courtship, in which the end of the abdomen is made more accessible
to the male by lowering both the end of the abdomen and the hindlegs.
(Courtesy of Otte, 1970, Misc. Publ. Mus. Zool., Univ. Michigan,
No. 141.)
Oviposition behavior differs among species of grasshoppers. Most
species lay their eggs in the ground. Females of some species choose
bare ground, while others choose to lay among the roots of grasses
or forbs. A female ready to lay often probes the soil several times
before finally depositing a clutch of eggs. Experimental evidence
indicates that probing is a means by which a female obtains sensory
information on the physical and chemical properties of the soil.
The ovipositor is supplied with a variety of sensilla that allow
a female to monitor soil conditions. Temperature of the soil must
be favorable, water content must be in a suitable range, and acidity
and salt content of soil must be tolerable. Females of some species
select loose soils such as sand, some rocky soils, while the majority
of females prefer compact loamy soils for oviposition. The pods
of different species vary in size, shape, and depth in the soil.
The latter condition profoundly affects incubation temperature,
hatching, and egg survival.
A female actively depositing eggs is often attended by one or more
males. Depending at least partly on clutch size, females take from
25 to 90 minutes to lay a full complement of eggs. After withdrawing
her ovipositor a female will take a minute or two to cover the aperture
of the hole with particles of soil and ground litter. The female
uses either her ovipositor or her hind tarsi, depending on species,
to do this. The act appears to be instinctive maternal care that
provides some protection for the eggs from predation by birds, rodents,
and insects. In certain species, as soon as the female retracts
her ovipositor, the attendant male mates with her; females and attending
males of other species merely walk away from each other.
Grasshoppers have different ways of avoiding excessively high temperatures
that may occur in summer for a few hours each day. When ground temperatures
rise above 120°F, individuals of certain species climb plants, some
to a height of only 2 inches on grass stems, others climb higher
(5-12 inches), and some even higher on tall vegetation such as sunflower
in the southern mixedgrass prairie or a tall cactus plant in the
desert prairie. In these positions individuals usually rest vertically
with the head up on the shady side of the plant. Many ground-dwelling
species first raise up on their legs or stilt, but as temperatures
rise further, they crawl into the shade of vegetation. Individuals
of other ground-dwelling species may stilt, then move away from
the hot bare ground and climb on top of a short grass such
as blue grama. They face the sun directly so that the least body
surface is exposed to the radiant rays. In this orientation the
grasshoppers are 1 to 2 inches above ground and rays of the sun
strike only the front of the head.
Characteristic behavioral responses regularly observed by grasshopper
scouts and collectors of insects are the jumping and flying of grasshoppers,
apparently to escape capture. The stimuli initiating these responses
may be of many kinds but circumstantial observation indicates that
movement of the collector's image across the compound eye is usually
the primary stimulus. One may noisily push a stick on the ground
close to an individual without eliciting a response. Also other
insects, such as large darkling beetles or large grasshoppers, may
crawl close and elicit no response until they touch a resting grasshopper.
Phytophilous species of grasshoppers may not jump or fly to escape
intruding scouts and insect collectors, but may shift their position
away from the intruder to the opposite side of a stem, retreat deeper
into the canopy, or drop to the ground. Collectors and scouts entering
the habitat of dense populations of grasshoppers may get the impression
of continuous movement by these insects, but investigation of their
time budgets indicates that during most daylight hours they are
largely quiescent (46 to 80% of the time). Considerably less time
is devoted to feeding, locomotion, mating, and oviposition.
The adults of most species of grasshoppers possess long, functional
wings that they use effectively to disperse, migrate, and evade
predators. Adults regularly fly out from deteriorating habitats
caused by drought or depletion of forage, but they may also leave
a site for other reasons. Some individuals of a population instinctively
fly and disperse while others remain in the habitat. The most notorious
migrating grasshoppers are Old World locusts, Schistocerca gregaria
and Locusta migratoria. Several species of North American
grasshoppers are likewise notable migrators: Camnula pellucida,
Dissosteira longipennis, Melanoplus sanguinipes, M. devastator,
M. rugglesi, Oedaleonotus enigma, and Trimerotropis pallidipennis.
Past direct observations of grasshoppers in their natural habitat
have revealed various behavioral responses among species. Each appears
to have its own way of reacting to a battery of environmental factors.
Because a relatively few species have been investigated, much remains
to be discovered. The whole subject of grasshopper behavior provides
a fertile area for research both in the field and in the laboratory.
Acquiring sufficient information on the various aspects of grasshopper
behavior will not only serve to improve integrated management of
pest species and the protection of beneficial ones, but will also
advance the science of animal behavior. Numerous observations of
the behavior of common western grasshoppers have been made during
the course of this study. Results of these observations are reported
later in the treatment of individual species.
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Introduction
| External Anatomy |
Names and Species | Grasshopper Populations
| Life Cycles | Seasonal
Occurrence and Behavior | Collections
and Survey | Grasslands of North America
| Food Plants | Glossary
| Selected References |
Species Fact Sheets | (PDF)
|