Insects" from Atlas of the Sand Hills

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October 1998

"Insects" from Atlas of the Sand Hills

Brett C. Ratcliffe University of Nebraska-Lincoln, [email protected]

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Ratcliffe, Brett C., ""Insects" from Atlas of the Sand Hills" (1998). Papers in Entomology. 44. https://digitalcommons.unl.edu/entomologypapers/44

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Brett C. Ratcliffe Curator of Entomology University of Nebraska State Museum

What is life? It is the flash of the firefly do the job nor adequate resources devoted sects in the Sand Hills tolerate drier and in the night. It is the breath of a buf- to such an undertaking. No one knows, windier conditions and greater solar ra- falo in the winter time. It is the little therefore, how many species of insects diation. They have also been successful shadow which runs across the grass occur in Nebraska, although the number in surviving the periodic fires that are so and loses itself in the sunset. undoubtedly reaches the thousands. necessary for maintaining native grass- However, a number of specific in- lands. In fact, the mosaic of habitats par- —Crowfoot, 1890, Blackfoot sect groups in Nebraska have been sur- tially created by fire has probably con- Confederacy Spokesman veyed, including in the Sand Hills. These tributed to increased insect diversity in range from simple checklists to exhaus- the Sand Hills. Introduction tive monographs; some are old and largely With the introduction of row crops outdated, while others are new and very grown as monocultures (growing a single More than a hundred years after usable. Unfortunately, it is possible in a crop in the same place many years in a Crowfoot's brief, but eloquent statement single chapter to give only a limited over- row), often with irrigation, a few insect about the nature of life, our knowledge view of the insects in the Sand Hills, not species that formerly occurred in the Sand of the insects of the Sand Hills, though so much because we still lack baseline Hills in relatively low numbers now oc- greatly advanced, is still so fragmentary data, but because we already have so casionally reach economically damaging that many species remain as only little much; volumes could be devoted to it. levels (Wedberg and others, 1975). On shadows running across the grass. Imag- Such is the nature of dealing with a group the other hand, some formerly success- ine. One out of every four living things of organisms numbering, perhaps, 10 mil- ful species have now disappeared from on Earth is a beetle! Add in all the re- lion species. native prairie because of the severe dis- maining insects, and it becomes clear that Because the Sand Hills are, for the turbance caused by agriculture. a biotic survey for all the insects of any most part, a treeless landscape of grass- given region is an immense undertaking. covered sand dunes, most of the insects In fact, a survey of all the insects in any are adapted to sandy habitats and a dry, Insect Natural History one state in the nation has never been suc- continental climate, as well as to the kinds cessfully completed because there are of plants and other animals that live there. Insects are, directly or indirectly, an simply not enough insect researchers to Unlike most forest-adapted species, in- important part of our everyday lives.

The endangered American burying beetle gets ready to go to work on a dead kangaroo rat (see sixth paragraph of section on Coleoptera). Honeybees (Apis mellifera L.) have become important pollinators of alfalfa and other fruit- and vegetable-producing plants, but they are not native to North America. Consider, for example, that one acre of red clover contains about 216 million individual flowers, and every one of these must be visited by an insect before Fig. IN-2. Minute springtails often occur in it will produce seed (Borror and others, huge numbers in moist soil, especially around 1989). Honeybees provide most of the Sand Hills lakes. pollinating necessary for these plants, thus greatly increasing seed yield. who wish to see vertebrates in Fig. IN-1. Though solitary, andrenid bees can Without insect pollinators, many of tomorrow's national parks need to be abundant in spring and often nest together the prairie plants would disappear and give serious consideration to the in the ground where vegetation is sparse. with them the animals that rely on them. place of insects in the food and com- And so would begin the collapse of most petition webs leading to those verte- Most people don't recognize the signifi- of the complex food chain where insects brates (Janzen, 1987). cance of the relationship between insects feed on other insects; where fish, birds, and humankind. Few realize that insects and mammals feed on insects and each Without insect scavengers to recycle are far more beneficial to our lives than other and so on. wastes, pastureland, rangeland, and prai- they are harmful. Why this misunder- The yucca moth (Tegeticula rie would be decimated as grasses became standing? It results mostly from media yuccasella [Riley], family Prodoxidae) is smothered by ever-increasing amounts of coverage of the more sensational, nega- the sole pollinator of the many yucca feces (solid waste) from livestock and tive aspects of insects (killer bees, crop plants in the Sand Hills. Both moth and wild animals. The sandy soils of the Sand pests, science fiction monsters, or "creepy plant rely on each other exclusively for Hills have been continually enriched by crawlies" in your home) and from a criti- their existence. Female moths insert their the action of dung beetles (Scarabaeidae), cal absence of education in our elemen- eggs into the ovary of the yucca flower other beetles, and flies that bury feces of tary and secondary schools about inver- and then force a wad of pollen they have animals. Aeration and enhancement of tebrates. Consequently, most Americans collected from other yuccas into the long, nutrients in the soil by dung beetles pro- are actually afraid of most insects and do funnel-shaped stigma (the very end of the motes the growth of forbs (broadleaf not appreciate their value economically, pistil typically sticking out of the center herbs), shrubs, and wind-pollinated ecologically, or aesthetically. of the flower and receiving the pollen). grasses. Bison and then cattle relied on Insects are incredibly important in The moth larvae eat some of the develop- these plants for food, food made possible, our lives. While it is true that just a very ing seeds that result from cross fertiliza- in part, by scavengers. All life on the prai- few can cause substantial losses to our tion, but enough seeds are left uneaten to rie is intricately interconnected. agriculture and structures and a few oth- ensure survival of the yucca. Insect predators and parasites are ex- ers transmit diseases to us and our live- Actually, two-thirds of all flowering tremely important in controlling many in- stock and crops, the vast majority are ei- plants depend on insects for cross polli- sect pests that would otherwise attack our ther directly or indirectly beneficial to hu- nation and, hence, survival. Insects and crops. Mantids and the reddish-orange mankind. In fact, without insects, the plants have co-evolved over millions of (spotted or not) ladybird beetles are of- Sand Hills prairie, with its associated years to develop these special, mutual re- ten mass-reared and deliberately released plants and animals, would not exist as we lationships. Many flowers have special to prey upon pest insects, while other know it. structural features to ensure that insects naturally occurring predators such as Insects are invaluable pollinators. In will pollinate them. We would do well to ground beetles and aphidlions (aphid fact, the pollinating services of insects are remember that the annual economic ben- predators) are working on our behalf. worth about $19 billion annually in the efits in the United States resulting from Many minute wasps and flies are valu- United States (Borror and others, 1989). just insect pollination is greater than the able parasites of aphids, cutworms, and Almost any plant on the prairie with con- losses from all kinds of insect damage weevil larvae that often cause economic spicuous flowers is insect pollinated, combined. damage to crops and fodder. The larvae whether it be by butterfly, moth, beetle, Without insects as food, most of our of parasitic insects develop inside the fly, or bee. Insects that prey upon polli- song birds and game fish would disappear. body of their host and consume it. This nators also become inadvertent pollina- relationship results in the death of the host tors themselves when they carry pollen Insects are major connectors between but not before the parasite completes its from one flower to the next while hunt- different species in their habitat, in- own life cycle. ing. Wild bees and bumblebees are the cluding species that have long been One of the principal reasons for the most important pollinators of prairie in the eye of the conservation com- biological success of insects is their rapid flowers in the Sand Hills. The large munity. By consuming large quanti- rate of reproduction. This helps them bumblebees are social insects (like hon- ties of plants and plant parts on the adapt to changing climates and habitats eybees), whereas wild bees are all soli- one hand, and by being consumed in through vast stretches of time. For ex- tary. Wild bees (sand bees, sweat bees, enormous amounts by vertebrates on ample, the greatest swarm of desert mi- leafcutter bees, alkali bees; fig. IN-1) are the other hand, insects in great part gratory grasshoppers ever recorded was ground-nesting, but their burrows may be determine the structural relationship in South Africa, where it covered an esti- so close together so as to form colonies. between larger organisms. Those ... mated area of 2,000 square miles! The swarm was blown out to sea, and when Wright (1970) suggested that the aquatic, and the many unpolluted ponds the tide washed back their bodies, they sand forming the dunes in the Sand Hills and streams in the Sand Hills support high formed a drift along the beach 50 miles was probably deposited by Pleistocene numbers of mayflies. The adults, being long and 4 feet high! A swarm of this size winds off of or near glaciers in an envi- relatively weak fliers and short-lived, are contains an estimated 250 billion hoppers ronment that was characterized by Smith most common near water. weighing about 500,000 tons. Springtails (1965) as first a desert, with large, trans- Mayfly nymphs (young) feed prima- (Order Collembola) (fig. IN-2) are very verse dunes built by northerly winds, and rily at night on algae and bottom debris. abundant but seldom noticed because of later as an area of sparse vegetation with They are easily recognized by the pres- their small size. These pale, minute in- linear dunes superimposed on the older ence of leaf-like gills along the sides of sects are so numerous that it has been cal- dunes by winds from the northwest. The the abdomen. Mayflies have simple meta- culated that the top 9 inches of soil in one origin of the sand was probably the poorly morphosis, that is, gradually become acre of grassland contains 230 million of consolidated sediments of Tertiary age in larger and look more like an adult, as op- them; that works out to be 5,000 per southwestern South Dakota and eastern posed to passing through a pupal stage square foot. Macrotermes, an East Afri- Wyoming or from river-deposited sedi- (during which they change from a larva can termite, has a queen that lays an egg ment derived from these rocks about to a winged adult). Nymphs usually need every two seconds... or 4,300 every day. 10,000 years ago (Watts and Wright, one or two years to develop, but the adults Termite queens may live 50 years! 1966). live for only a day or two. Macrotermes is also the largest termite in The glaciations of the Pleistocene, Huge emergences of mayflies from the world; queens reach a length of 5 followed by gradual formation of the lakes and streams have occasionally been inches (Nebraska's termite queens are Sand Hills, affected insects living in Ne- recorded. Burks (1953) observed piles of about one-half inch). Fruit flies are small braska. During glacial times, much of the mayflies 3 feet deep in Illinois, which insects about the size of a pinhead. They insect fauna in the state retreated to the caused serious traffic problems on the af- can have 25 generations a year, and each south, west, or east of the ice sheets. Fol- fected roads. Both adult and nymphal female lays about 100 eggs. A single pair lowing the withdrawal of glacial ice and mayflies are an important food for many of these short-lived, tiny flies and their boreal forests, Nebraska was recolonized species of fish, as well as for birds, am- progeny could, if none died, increase in a from the south, west, and east by an in- phibians, and other insects. As such, they year to about 1041 flies. If you packed sect fauna characterized by taxa (types of are very beneficial insects. them together, a thousand to a cubic inch, insects) that had not occurred here in the they would form a ball of flies 96 million first place, as well as by those whose an- miles in diameter . . . the same distance cestors had previously been driven out by as from the Earth to the Sun! glaciers. Because of this, the insect fauna of the state is unique and is a recent mix Fig. IN-3. Adult mayflies usually live for only of both eastern, western, northern and a day, while the aquatic nymphs live for one Biogeography southern faunas that have converged here to two years. and are near the limits of their respective Contrary to the stories about Mid- ranges (Ratcliffe, 1988a, 1991). An insect western prairie states having an impov- fauna unique to the Sand Hills or, for that erished number of species, Nebraska's matter, Nebraska does not exist, although Sand Hills have a diverse insect fauna that a few native or isolated species may oc- reflects the multitude of habitats that occasionally be found there. Kaul (1986) cur there. Most insect species are special- observed that the flora was also of exter- ized in their microhabitat preferences, and nal origin with nearly all of the plant spe- so upland dunes support different species cies having colonized from elsewhere, than moist canyons, wetlands or forest especially from the southeast and south- edges. west. Many eastern species of both ani- Near the end of the Pleistocene (Ice mals and plants extend west to approxi- Age) glaciations, much of Nebraska was mately 100 degrees west longitude in covered by forests of spruce and pine Nebraska—about the middle of the (Picea spp. and Pinus spp.) and open state—and many western species dis- parkland that grew in a boreal (cool, persed eastward to roughly that same di- moist, northem-like) climate. Pollen data, viding line. as well as fossils of late Pleistocene forest mammals (musk-ox, stagmoose, and jaguar), indicate the presence of spruce The Insects tundra in north-central Nebraska (Schultz and others, 1985). About 12,000 years Ephemeroptera ago, as the climate became warmer and drier after the last glaciation, following Mayflies (fig. IN-3) are small to me- the late glacial dominance of boreal for- dium-sized, elongated, soft-bodied insects ests, prairie grasslands developed and with two or three long, threadlike tails. spread throughout the region that included The adults have delicate, membranous Nebraska. These grasslands are, therefore, wings, and the forewing is usually large relatively young. and triangular. The immature stages are the prairie grasses, but few roaches (Or- der Blattaria) live in the Sand Hills because it is too dry. Most species of Orthoptera are plant feeders, and some are important pests of cultivated plants and rangeland grasses. The crickets (Gryllidae) and long-horned grasshoppers (Tettigoniidae), or katydids are remarkable for the songs they produce by stridulation, or rubbing one body part against another. Different species have different songs, and it is the males that sing to attract females. Different species can actually be distinguished in the field based on their unique songs. Grasslands are the home to a wide array of Orthoptera. Sand Hills prairie is no exception, and many species of short- horned grasshoppers (Acrididae) occur there (Hagen, 1970; Joern, 1982, 1983; Walking sticks are often abundant on prairie grasses. Hagen and Rabe, 1991). Orthoptera, in the broad sense, have long been studied Odonata its prey by scooping and tumbling it into in Nebraska because of their economic the basket of legs. importance, and several state lists have Heavy-bodied dragonflies and their Dragonflies and damselflies are most been published (Bruner, 1893; Hauke, smaller and less agile relatives, damself- active during the sunny part of the day, 1949, 1953). lies, frequent the shores of ponds, but swarms of dragonflies can often be marshes, and streams. About 80 species seen at dusk hunting for the midges and are found in Nebraska (Pruess, 1967). The mosquitoes that are starting to fly. Al- young of Odonata, called naiads, are though fearsome predators to other small aquatic and are voracious predators on insects, dragonflies are totally harmless just about anything they can subdue, in- to humans (Ratcliffe, 1991c). cluding tadpoles, snails, and small fish. Most adult dragonflies feed primarily on Orthoptera small, flying insects such as mosquitoes, midges, and small moths. Larger dragon- The Order Orthoptera consists of flies will capture bees, butterflies, and grasshoppers, crickets, and katydids. even other dragonflies. Adults catch their Walking sticks, mantids, and roaches used prey while flying, using a basket-like ar- to be included here, but the current clas- Stink bugs are not really as unpleasant as their rangement of the legs, which are useless sification places each of those groups in name suggests. for walking, but serve much like an aerial a separate order of its own. Walking sticks net. In full flight, the dragonfly captures (Order Phasmida) are often abundant on The short-horned grasshoppers (Acrididae) are well known to everyone Long-homed grasshopper (katydid). Males are noted songsters in late summer. because of their conspicuous size and abundance in fields and along roadsides. Often called locusts, most acridids are brown or gray in color, and some have brightly colored hindwings. Some parts of Nebraska were devastated in the 1870s by huge, migratory swarms of the Rocky Mountain locust (Melanoplus spretus Walsh) that consumed nearly every plant in their way. Crops disappeared, trees were stripped of leaves, and the ground moved like a crawling carpet. A single swarm observed during this time was estimated to contain 124 billion grasshoppers! Remarkably, this species is now extinct, the last specimen having been collected in 1902 (Lockwood and others, 1990). Even today, populations of some species in other parts of the world occasionally, though briefly, reach levels high hairlike antennae and, in the females, by the laterally compressed, blade-like ovipositor, which is used to deposit eggs. Most species are noted songsters, and they can especially be heard at night in the late summer in the prairie grasses. Their short chirps, buzzes, lisps and clicks are dis- tinct and, once identified, are rarely con- fused with the pulsating and rhythmic chirping of field crickets or tree crickets. The crickets (Gryllidae) also have long antennae but differ from the tettigoniids (see above) in having the front wings bent down sharply at the sides and by the presence, in the females, of a needle-like ovipositor. Most species are well-known songsters and are active at night. Adults reach their highest numbers in late summer. They are omnivorous and feed ravenously on plants and dead insects, especially grasshoppers. Some species of leafhoppers transmit plant viruses to certain crops. Hemiptera and Homoptera Most Hemiptera and Homoptera The so-called bugs (Order Hemi- feed on plant juices, using their sucking ptera) and cicadas, aphids, and leafhop- mouthparts although some bugs are pre- Fig. IN-4. Chinch bugs are sometimes serious pers (Order Homoptera) are well repre- daceous on other insects. For example, pests of small grains. sented in the Sand Hills. Aside from the some giant water bugs (Belostomatidae) works of Zimmer (1911) on the will feed on tadpoles and small fish, and enough to cause economic damage to Pentatomoidea (stink bugs and their rela- swallow bugs (Cimicidae: Oeciacus rangeland and crops (Capinera and Th- tives) and Williams (1910) on the vicarius Horvath) are blood-feeding, ex- ompson, 1987), but swarms like those Aphididae (aphids), relatively little has ternal parasites of cliff swallows in the seen in the 1870s have not been seen since been written about the Nebraska fauna Sand Hills. in Nebraska. other than accounts of economically im- A number of species are aquatic. Nebraska has more than 100 species portant leafhoppers or aphids. Water boatmen (Corixidae), back-swim- of grasshoppers (see last page of this mers (Notonectidae), and water striders chapter), but most of the damage done to (Gerridae) are abundant in the ponds and crops and rangeland here is caused by four Fig. IN-5. Dog-day cicadas provide one of the marshes in the Sand Hills, as well as in species of acridids: the migratory grass- most typical sounds of hot, summer days in stock tanks. Chinch bugs (Lygaeidae: hopper, Melanoplus sanguinipes (Fabr.), the Sand Hills. Blissus leucopterus Say) (fig. IN-4) were the differential grasshopper, M. of economic importance historically in the differentialis (Thomas), the two-striped Sand Hills because they attacked wheat, grasshopper, M. bivittatus (Say), and the oats, sorghum, and corn. Entire fields of red-legged grasshopper, M'. femurrubrum grain were destroyed when these bugs (DeGeer) (Borror and others, 1989). All were abundant. of these species occur in the Sand Hills. Among the Homoptera, the cicadas Pfadt (1988 and ongoing) is producing (Cicadidae) (fig. IN-5) are probably the loose-leaf fact sheets with color illustra- most easily noticed in the Sand Hills, but tions on the grasshoppers of the western they are more usually heard than seen. United States. These guides enable iden- Male cicadas produce a courtship song tification and discuss life histories. characteristic of their own species; fe- The largest grasshopper in the state males, however, are incapable of produc- occurs in the Sand Hills. It is the lubber ing sound. Often incorrectly called lo- grasshopper (Brachystola magna Girard), custs, cicadas normally live in forested a flightless species that reaches a length areas where they lay eggs on trees and of 2.4 inches (6 cm). The rainbow grass- the nymphs feed on tree roots. In grasshopper (Dactylotum pictum Thomas) is lands such as the Sand Hills, they have another, smaller flightless species whose had to adapt to the absence of trees. One body is beautifully marked with bands of > of the ways they have done this is by hav- black, red, yellow, and pale blue. y ing a shorter life cycle, that is, four or five The long-horned grasshoppers e years long instead of the customary 10 to (Tettigoniidae) are distinguished from the 17 years! In the Sand Hills, yucca is fre- short-homed grasshoppers by their long, quently the tallest plant around and so Green lacewing adult. The larvae are beneficial predators on aphids.

replaces trees as a host for grassland ci- habitat and possibly to escape the keen such aggressive predators that they might cadas. eyes of predators. The spittle is a mixture eat each other. Lacewing larvae, called Leafhoppers (Cicadellidae) are of fluid voided from the anus and a mu- aphidlions, are beneficial because they are abundant and feed on the variety of cous-like substance secreted by glands on among the primary predators of aphids. grasses in the Sand Hills. Some species the abdomen. The two fluids are mixed Adult lacewings are often attracted to reach economic thresholds when they mi- into a froth by the rapid movement of the porch lights at night. Look closely at a grate into fields of cultivated crops. Many hind legs. These masses of spittle are fre- live adult and you will see that it has beau- species are carriers of plant diseases and quently seen on the stems of grasses. tiful golden-colored eyes. so are of economic importance. Spittle- Antlions (Myrmeleontidae) are com- bugs (Cercopidae) resemble leafhoppers Neuroptera mon in the Sand Hills, and the telltale, but are brown or gray in color and hop. conical pits made by the larvae are found They feed on grasses and herbaceous Adult neuropterans are soft-bodied, in the fine sand under nearly every rock (broadleaf) vegetation. The nymphs sur- membranous-winged insects. Most of the ledge or overhang. The larvae are large- round themselves with a frothy, spittle- larvae are predaceous and are either jawed predators, most of whom lie in wait like mass in order to maintain a moist aquatic or terrestrial. The order is com- concealed in the bottom of their conical prised oflacewings, fishflies, dobsonflies, pits waiting for some hapless victim to Fig. IN-6. Adult dobsonflies are sometimes and antlions. Many species prefer drier tumble in. The adults are delicate, mem- attracted to lights at night. habitats, and so the Sand Hills is an ideal branous-winged insects that some might habitat for them. confuse with damselflies (Odonata); how- The dobsonflies and fishflies ever, antlions have knobbed antennae and (Corydalidae) have aquatic larvae, and the wings with many veins, whereas damsel- adults are generally found near water. The larvae are often called hellgrammites and are a favorite bait of fishermen. Dobson- A Colorado potato beetle is a significant pest flies are large insects and have front wings of some crops. 2 inches (5 cm) or more in length. Male dobsonflies (fig. IN-6) appear ferocious with their hugely elongated mandibles (jaws), but they are harmless. Females, with their shorter and more powerful mandibles, could provide a painful nip if handled carelessly. Green lacewings (Chrysopidae) are common insects, with 87 species in North America. Each egg is laid on the end of a long stalk, possibly because the larvae are and they can emit a foul-smelling fluid that discourages enemies wanting to eat them. "Chemical warfare" is highly developed in many species and is used with great success in defense. The phytophagous leaf beetles (Chrysomelidae) are very rich in species in the Sand Hills. Most are brightly colored and feed on foliage. Some species of leaf beetles (flea beetles, corn root- A weevil, one of many hundreds of species in worms, Colorado potato beetle, and the Nebraska spotted cucumber beetle) are pests of cultivated plants. Weevils (Curculionidae) are also abundant in the Sand Hills but flies have small, slender antennae and are not as easily seen because most are wings with only a few veins. There are gray, brown, or dull-colored. Weevils are 89 species in North America, and most easily recognized by their elongated prefer dry, sandy areas. snouts. All of the 2,600 species in North America are plant feeders, and many spe- Coleoptera cies are serious pests in other parts of the country. Aside from billbugs in turfgrass, One out of every four living things weevils do not seem to reach economic on Earth is a beetle! In numbers, that thresholds in the dry Sand Hills. translates to about 450,000 species. There Tiger beetles (Cicindelidae) are are, for example, more kinds of weevils brightly colored and a favorite with col- Fig. IN-7. Ground beetles, such as this (one of the larger families of beetles) than lectors. They run and fly rapidly and are Bembidion sp., are often found in marshy all mammals, birds, reptiles, and amphib- very difficult to approach because they areas around streams and lakes. ians in the world combined! The beetle are so wary. They frequent sandy habi- fauna of the Sand Hills is rich and varied, tats and shorelines where they are fero- Gothenburg, both along the Platte Valley but even today no one knows how many cious predators on smaller insects. With and in the uplands. Formerly occurring species exist there because so few stud- 32 species, Nebraska has one of the high- in the eastern half of North America, this ies have been done in Nebraska. With est number of species of any state, and species is now found in only six states. about 125 families of beetles, it is impos- 12 species occur in the Sand Hills alone. Nebraska has one of the highest popula- sible to do them justice in this format. Nebraska's tiger beetles were reviewed by tions of this rare insect. Aside from sen- Hundreds of species occur in the Sand Carter (1989), and Ratcliffe (1990b) sitivity to human disturbance, no one Hills, but only a very few are mentioned briefly summarized their life history. knows why its numbers have declined so here. The huge family of ground beetles precipitously during the last half century. Many kinds of water beetles (Carabidae) (fig. IN-7), numbering some (Dytiscidae, Hydrophilidae, Gyrinidae) 2,500 species in North America, has not inhabit the ponds, lakes, and stock tanks been studied in Nebraska, although in the Sand Hills. These bullet-shaped, Clopton (1991) provided a synopsis of the Fig. IN-8. Dung beetles are very beneficial amphibious beetles feed on algae, decay- tribe Scaritini. Most ground beetles are insects that help to clear pastures of grass- ing vegetation, and small aquatic inver- predators, and hence beneficial, while a smothering dung pats. tebrates. Their larvae, however, are vora- few feed on seeds. Most are nocturnal, cious predators that feed on worms, and many are attracted to lights at night, leeches, tadpoles, other insect larvae, and often in large numbers. even small fish. Click beetles (Elateridae), Ratcliffe (1996) provided a detailed so called because of their ability to right faunal survey of the carrion beetles themselves by snapping their head and (Silphidae) of the state. Carrion beetles thorax down onto the substrate so hard are an important part of a vast host of that they make an audible "click" while scavengers that break down and recycle launching themselves upward and over, elements found within dead animals. Car- are abundant. The larvae (wireworms) of rion beetles belonging to the genus some click beetles are destructive because Nicrophorus actually bury small, dead they feed on seeds and roots. Ladybird animals (hence the name sexton beetles, beetles (Coccinellidae) and luminescent referring to the old church grave digger) fireflies (Lampyridae) are well known to in order to provide their young with food. everyone. The black darkling beetles The rare American burying beetle (Tenebrionidae) are abundant in the Sand (Nicrophorus americanus Olivier) (see Hills. When disturbed, members of the first page of this chapter) is an endan- genus Eleodes seem to nearly stand on gered species that is found in the Sand their head as they run. While perhaps Hills in Cherry and Keya Paha counties, amusing to us, this is a defensive posture, as well as between North Platte and beetle in the Sand Hills is the cottonwood borer (Plectrodera scalatorF'dbr.), which is 1.6 inches (4 cm) long and strikingly marked in enamel black and white. Adults can be found around cottonwood trees in which they breed, and they are also attracted to lights. Attractive red and black longhorn beetles belonging to the genus Tetraopes are fairly common on the flowers of milkweeds. Nebraska's longhom beetles were summarized by Baker (1977). Diptera

The true flies, as opposed to dragonflies, caddisflies, butterflies, etc., are characterized by having only one pair of wings. The second pair of wings have been reduced to small, knob-like struc- Swallowtail butterflies, so-called because of the tails on the hindwings, are the largest butterflies tures called halteres that are used to main- in Nebraska. tain balance while in flight. Flies belong Some of the most beautiful and larg- mals only. Most dung beetle behavior and to one of the largest orders of insects, and est insects on Earth are scarab beetles nest building is adapted to grassland con- they are found nearly everywhere using (Scarabaeidae). Of all the beetles, scar- ditions, which is not too surprising since almost every kind of habitat. Many spe- abs are among the best known because of most large mammal herbivores (plant eat- cies have predatory or herbivorous their beauty, fascinating life histories, and ers) occur in this habitat. Some scarab spe- aquatic larvae, and the many Sand Hills economic importance. There are about cies form balls of dung and roll them away lakes, wetlands and streams support a di- 1,500 species in North America, and Ne- to escape the intense competition for a verse array of mosquitoes (Culicidae), braska has nearly 200 species. Approxi- limited food (the dung) at a given dung midges (Chironomidae), black flies mately 13 percent of Nebraska's scarabs source. After finding a suitable place, the (Simuliidae), soldier flies are found in the Sand Hills. A few spe- dung ball will be buried and either con- (Stratiomyidae), horse flies (fig. IN-9) cies {Polyphylla spp., Bolboceras spp.) sumed or used as a brood ball for a de- and deer flies (Tabanidae), among others. are found only in sandy habitats veloping larva. The larvae of these flies often constitute (Ratcliffe, 1990a), while many others are Their benefit to maintaining the an important food item for many fresh- adventive. Several small species are health of grazing lands while, at the same water fish. The females of mosquitoes, found only in the burrows of prairie dogs time, reducing the breeding spots for horse flies, and deer flies are blood feed- and gophers and are rarely encountered many pest species of flies, should not be ers, primarily on warm-blooded verte- (Ratcliffe, 1988b). One of the most beau- underestimated. As far as food for dung brates. Tate and Gates (1944) summarized tiful scarabs in the Sand Hills is Phanaeus beetles is concerned, the introduction of the mosquitoes of Nebraska. vindex Macleay, a horned dung beetle cattle into the Sand Hills effectively re- The largest fly in Nebraska occurs with shining metallic green wing covers placed the bison that formerly occurred only in the Sand Hills. It is a robber fly and a copper-colored thorax. The several there. Ratcliffe (199 la) provided a com- (Asilidae: Proctacanthus milberti species of dung beetles (fig. IN-8) that plete summary for the state's scarab beetle occur in the Sand Hills are critical to the fauna. Fig. IN-9. Female horseflies are bloodsucking recycling of feces. These beetles bury Stag beetles, or pinch bugs and may become serious pests of livestock and dung beneath the ground, and this serves (Lucanidae), are more common in east- people. The males feed on pollen and nectar. as food for their larvae. Most dung beetles ern Nebraska, but one large species are opportunistic and feed on the excre- (Lucanus placidus Say) is found in the ment of many kinds of animals, but some Sand Hills (Ratcliffe, 1991b). Adults are are specific for the feces of certain ani- attracted to lights, and they look intimi- dating because of their greatly enlarged Robber flies are aerial predators, taking any mandibles. Appearances aside, they are, small insect they can subdue. in fact, relatively harmless, although the mandibles could pinch if one placed a fin- ger between them. Long antennae characterize most of the longhom beetles (Cerambycidae). The adults feed on pollen, leaves, and wood, while the larvae bore into wood and roots. The family is popular among collectors because many species are large and brightly colored. The largest longhorn be found hovering about flowers, upon which they feed, often being mistaken for bees. A synopsis of the syrphids of Ne- braska was given by Wehr (1922). The state of our knowledge about the pomace flies (Drosophilidae) in Nebraska is probably typical for most families of Diptera in the state. These tiny flies (0.12- 0.16 inches [3-4 mm]) are seldom col- The larvae of monarch butterflies feed on lected and infrequently studied. As a con- milkweed. sequence, our knowledge of them is ex- Typhiid wasps are parasitic on white grubs in ceedingly fragmentary and, aside from the subirrigated meadows. Macquart) and may reach a length of 1.6 list produced by Williams and Miller inches (4 cm), a truly large fly! Robber (1952), we know little about the species flies frequent open, sunny areas looking composition in Nebraska, let alone their (Prunus virginiana L.) grows. Ratcliffe for bees, butterflies, tiger beetles, wasps, natural history and interactions with other (1992) briefly summarized their life his- or grasshoppers. When they find suitable plants and animals. In spite of their small tory. The monarch butterfly (Danaus prey, they grasp it, inject digestive juices size, however, they and all other insects plexippus L.), which adapts to nearly ev- into it with their syringe-like beak, and are essential to nature's ecological bal- ery environment, takes nectar from a va- then suck out the body fluids. Their lar- ance. riety of flowers while their larvae are re- vae, equally voracious, feed on grasshop- stricted to feeding on milkweeds. Adult per eggs and beetle grubs in the soil. Lepidoptera monarchs have been known to fly as far The blow flies (Calliphoridae) and as 2,000 miles on their annual migration flesh flies (Sarcophagidae) are primarily Butterflies and moths have a lower south to overwinter in Mexico. Not only scavengers, and the larvae feed on car- diversity in the Sand Hills relative to the is the distance incredible and the route ex- rion and excrement, thus providing a rest of the state. There are many reasons ceedingly complex for a tiny animal, but valuable service by cleaning up the land- for this distribution, but principal among think about how many streets and high- scape. On the other hand, face flies, stable them is the predominance of grasses, in- ways had to be successfully crossed with- flies, and horn flies (Muscidae: Musca stead of broadleaf plants upon which out getting hit by rushing, multi-ton ve- autumnalis DeGeer, Stomoxys calcitrons Lepidoptera larvae usually feed. Adult hicles. Red admirals (Vanessa atalanta L.) L., and Haematobia irritans L., respec- butterflies are generalist flower feeders and painted ladies (V. cardui L.) are abun- tively) are biting flies and can be signifi- except for those few that do not feed. dant on thistles and catnip. cant pests of livestock in the Sand Hills. Swallowtail butterflies (Papilio glaucus The giant silk moths in the family The Syrphidae (flower flies) and L. and P. multicaudata Kirby, family Saturniidae (Antheraea polyphemus Bombyliidae (bee flies) are often abun- Papilionidae) are restricted to riparian Cramer, the Polyphemus moth, and dant in the Sand Hills, where they may (river) canyons where chokecherry Hyalaphora cecropia L., the Cecropia

The gray hairstreak takes nectar from flowering dogbane. meadows or marshes where dock (Rumex spp.) grows, and a few hairstreaks (also Lycaenidae) keep close to marshes or wet meadows. The butterfly and moth species of the Sand Hills show more affinities with western taxa, although some eastern species occur there also. With good flight capa- bilities, a number of migrant Lepidoptera are also periodically found in the Sand Hills, but they do not breed there. Anno- tated lists of butterflies of Nebraska were provided by Barber (1894), Leussler (1938, 1939), and Johnson (1973). Dankert and others (1993) produced but- Fig. IN-10. Female cicada killers are the terfly distribution maps for Nebraska. largests wasps in Nebraska. Hymenoptera moth) have, on rare occasions, been taken in the Sand Hills, where their larvae feed The sawflies, ants, wasps, and bees on willows (Salix spp.) or cottonwoods represent the most beneficial order of in- (Populus spp.) (Ratcliffe, 1993) found in sects. Many species are valued because blowouts or along water courses. The at- they are parasites or predators of insect tractive, black and white Buck Moth pests or are important pollinators of Fig. IN-12. All the ants usually seen in a nest (Hemileuca nevadensis Stretch, also a sat- plants. The members of the Order Hy- are females. Winged males appear only for a urniid) is most common in the Sand Hills, menoptera have fascinating life histories brief time during the year. where the larvae feed on willows and pos- and show a great diversity of habits and sibly cottonwoods. The stout-bodied complexity of behavior, culminating in sphinx moths (also known as humming- the social organization of wasps, bees, and and Mickel (1917) provided a summary bird moths) have long tongues with which ants. of the Sphecoidea. they acquire the nectar of flowers over The relatively warm, dry, summer Some of the most conspicuous and which they so expertly hover. climate of the Sand Hills is conducive to certainly the largest, hymenopterans in the The Giant Skipper (Megathymus bees and wasps, and many species occur Sand Hills are the several species of streckeri Skinner) (Megathymidae) is re- there. In spite of a long tradition of spe- bumblebees (Bombus spp., family stricted in Nebraska to the Sand Hills but cialists studying Hymenoptera in Ne- Apidae) and the cicada killer (Sphecius is not native to them. Larvae feed on the braska, there is a lack of published sum- speciosus Drury, family Sphecidae). roots of Yucca glauca Nuttall, while the maries of this group in the state. Swenk Bumblebees prefer the nectar of red clo- adults, relying on huge amounts of stored (1907), Swenk and Cockerell (1907a, b), ver (Trifolium pratense L.) and also con- fat obtained in the larval stage, do not feed and LaBerge and Webb (1962) published tribute to alfalfa (Medicago sativa L.) pol- at all. Many blues (Lycaenidae) occur in short papers on selected groups of bees, lination (Ratcliffe, 1994). According to open Sand Hills prairie, where the larvae LaBerge and Webb (1962), various spe- feed on various legumes (Fabaceae), Fig. IN-11. Female velvet ants (actually a cies of Bombus are probably the princi- while adult hairstreaks and coppers (also wasp) have a powerful sting. pal pollinators of various prairie plants in Lycaenidae) take nectar from flowering Nebraska, such as prairie clover (Dalea dogbane (Apocynum spp.). spp.), milk-vetch (Astragalus spp.), thistle The numerous marshes in the Sand (Cirsium spp.), mint (Monarda spp.), Hills provide islands of moist habitat for sweet clover (Melilotus spp.), and sun- several butterfly species. Fritillaries, or flowers (Helianthus spp.). silverspots (Speyeria and Boloria spp., Cicada killers (fig. IN-10) are large family Nymphalidae), use violets (Viola (1.6 inches [4 cm]) wasps conspicuously spp.) for larval food plants and may be banded in yellow and black. A female will found in marshy areas that support vio- seek out cicadas, which they paralyze (but lets. The Smokey Eyed Brown (Satyrodes do not kill) with their formidable sting. eurydice fumosus Leussler, family Alternately flying with it and then drag- Satyridae) is found in scattered colonies ging it, they will transport the cicada back in permanent marshes in the Sand Hills to a previously excavated burrow. The where the larvae feed on sedges (Carex helpless cicada is placed in the burrow; spp.). This butterfly has a spotty distribu- the wasp lays a single egg on it, seals the tion elsewhere in Nebraska and is now burrow, and leaves. The wasp larva, upon extinct in many parts of its former range, g hatching from the egg, finds an immobi- especially in the east. Some coppers, such .a lized storehouse of cicada food upon as Lycaena xanthoides dione Scudder^ which to feed. (Lycaenidae), are restricted to prairie By virtue of their larger size and day- cause occasional economic loss. Consid- ment of Entomology Report No. 16, time activities, velvet ants (family ering that more kinds of insects occur pages unnumbered. Mutillidae) (fig. IN-11) are commonly there than all other Sand Hills plants and Hauke, H., 1949, An annotated list of the seen, especially in the late morning or late animals combined, we would do well to Orthoptera of Nebraska, Part I—The afternoon, when they may be viewed scur- increase our understanding of them and Blattidae, Mantidae and Phasmidae: rying about on the ground. Contrary to the role they play in the Sand Hills eco- Bulletin of the University of Ne- their common name, velvet ants are not system. braska State Museum, v. 3, p. 63-75. ants at all but female wasps that are wing- Hauke, H., 1953, An annotated list of the less. This condition, and the conspicuous Orthoptera of Nebraska, Part II—The short downy hair of many, have given rise References Tettigidae (sic) and Acrididae: Bul- to the common name, velvet ant. Although letin of the University of Nebraska most numerous in the southwestern Baker, J.R., 1977, A synopsis of the State Museum, v. 3, p. 1-79. United States, many are found in the Great Cerambycidae of Nebraska: Univer- Janzen, D., 1987, Insect diversity of a Plains. Nine genera are found in Ne- sity of Nebraska-Lincoln, Unpub- Costa Rican dry forest: why keep it, braska, and seven of these occur in the lished master's thesis, 262 p., plates and how?: Biological Journal of the Sand Hills. Most mutillids are parasitic 1-18. Linnean Society of London, v. 30, p. on other bees and wasps that also nest on Barber, H. G., 1894, A list of Nebraska 343-356. the ground in open, sandy areas. Rela- butterflies: The Nebraska Academy Joem, A., 1982, Distributions, densities, tively permanent dunes, devoid of most of Sciences, Proceedings, v. 4, p. 16- and relative abundances of grasshop- vegetation, are preferred nesting areas for 22. pers (Orthoptera: Acrididae) in a many mutillids, and so the Sand Hills rep- Borror, D.J., Triplehorn, C.A., and Nebraska Sandhills Prairie: Prairie resent an ideal habitat for them. About a Johnson, N.R, 1989, An Introduction Naturalist, v. 14, p. 37-45. dozen species of Dasymutilla are found to the Study of Insects, sixth edition: Joem, A., 1983, Host plant utilization by in the Sand Hills. A wide-ranging, large, Philadelphia, Saunders College Pub- grasshoppers (Orthoptera: Acrididae) red and black species, D. occidentalis L., lishing. from a Sandhills prairie: Journal of bears the common name of cow-killer be- Bruner, L., 1893, A list of Nebraska Or- Range Management, v. 36, p. 793- cause its sting is so painful. thoptera. Publications of the Ne- 797. True ants (Pormicidae) (fig. IN-12) braska Academy of Sciences, v. 3, p. Johnson, K., 1973, The butterflies of Ne- are widely distributed and are well known 19-33. braska: Journal of Research on the to almost everyone. They occur in nearly Burks, B.D., 1953, The mayflies or Lepidoptera, v. 11, p. 1-64. every terrestrial habitat and outnumber (in Ephemeroptera of Illinois: Illinois Kaul, R.B., 1986, Physical and floristic individuals) most other terrestrial animals. Natural History Survey Bulletin, v. characteristics of the Great Plains, in Ant colonies vary greatly in size, ranging 26, p. 1-216. Barkley, R.L., ed.. Flora of the Great from a dozen individuals to many thou- Capinera, J.L., and Thompson, D.C., Plains, p. 7-10: Lawrence, University sands. Different species nest in plant 1987, Dynamics and structure of of Kansas Press. stems, insect galls, decaying wood, or in grasshopper assemblages in short- LaBerge, W.E., and Webb, M.C., 1962, the ground. Most Sand Hills species nest grass prairie: Canadian Entomolo- The bumblebees of Nebraska: Uni- in the ground, where their galleries may gist, v. 119, p. 567-575. versity of Nebraska Agricultural Ex- extend from a few inches to many feet Carter, M.R., 1989, The biology and ecol- periment Station Research Bulletin underground. Some species in the genus ogy of the tiger beetles (Coleoptera: No. 205, p. 1-38. Formica (relatively common in the Sand Cicindelidae) of Nebraska: Transac- Leussler, R.A., 1938, An annotated list of Hills) build nests with a surface mound tions of the Nebraska Academy of the butterflies of Nebraska, with the of thatch made of dried grass stems and Sciences, v. 17, p. 1-18. description of a new species (Lepid.: leaves; the mound can be as high as 1-2 Clopton, R.E., 1991, A review of the Rhopalocera): Entomological News, feet (30-60 cm). Ants have varied feed- scaritinid beetles (Coleoptera: v. 49, p. 3-9, 76-80, 213-218, 275- ing habits and, depending on the species, Carabidae: Scaritini) of Nebraska: 280. consume fungi, plants, seeds, nectar, sap, Transactions of the Nebraska Acad- Leussler, R.A., 1939, An annotated list of honeydew, dead animals, and smaller liv- emy of Sciences, v. 18, p. 53-65. the butterflies of Nebraska, with the ing animals. Dankert, N., Nagel, H., and Nightengale, description of a new species (Lepid.: T., 1993, Butterfly distribution maps: Rhopalocera): Entomological News, Keamey, University of Nebraska at v. 50, p. 34-39. Summary Keamey, Department of Biology, p. Lockwood, J.A., Nunamaker, R.A., Pfadt, 1-34. R.E., and DeBrey, L.D., 1990, Grass- The insect fauna of the Nebraska Hagen, A., 1970, An annotated list of hopper glacier: American Entomolo- Sand Hills is richly diverse but still re- grasshoppers (Orthoptera, Acrididae) gist, v. 36, p. 19-27. mains incompletely documented. Imma- from the eleven panhandle counties Mickel, C.E., 1917, A synopsis of the ture stages and life-history information in of Nebraska: University of Nebraska Sphecoidea of Nebraska (Hy- particular are lacking. The insects living Agricultural Experiment Station Re- menoptera): University Studies, v. in the Sand Hills show adaptations for liv- search Bulletin No. 238, p. 1-60. 17, p. 342-456. ing in an environment largely made up of Hagen, A.R, and Rabe, G.C., 1991, Dis- Pfadt, R.E., 1988, Field guide to common grasses and with a climate characterized tribution maps of grasshopper spe- western grasshoppers: USDA by aridity. Many species are beneficial to cies in Nebraska based on three stud- APHIS/Wyoming Agricultural Ex- humankind, while only a few species ies: University of Nebraska Depart- periment Station Bulletin 912, sec- tions independently numbered. Ratcliffe, B.C., 1993, Nebraska's giant Watts, W.A., and Wright, Jr., H.E., 1966, Pruess, N.C., 1967, Checklist of Nebraska silk moths: NEBRASKAland Maga- Late-Wisconsin pollen and seed Odonata: Entomological Society of zine, v. 71,p.22-29. analysis from the Nebraska America, v. 22, p. 112. Ratcliffe, B.C., 1994, Bumblebees: Sandhills: Ecology, v. 47, p. 202-210. Ratcliffe, B.C., 1988a, Scarab beetles- NEBRASKAland Magazine, v. 72, p. Wedberg, J.L., Campbell, J.B., and Jeweled pollinators, dung feeders, 58-59. Helms, T.J., 1975, Bionomics of in- and horned giants. NEBRASKAland Ratcliffe, B.C., 1996, The carrion beetles sects associated with corn in the Ne- Magazine, v. 66, p. 30-37. (Coleoptera: Silphidae) of Nebraska: braska Sandhills: University of Ne- Ratcliffe, B.C., 1988b, A new species of Bulletin of the University of Ne- braska Agricultural Experiment Sta- Aphodius (Coleoptera: Scarabaeidae: braska State Museum, v. 13, p. 1-100. tion Research Bulletin No. 274, p. 1- Aphodiinae) from Nebraska: Trans- Schultz, C.B., Martin, L.D., and Schultz, 30. actions of the Nebraska Academy of M.R., 1985, A Pleistocene jaguar Wehr, E.E., 1922, A synopsis of the Sciences, v. 16, p. 87-89. from north-central Nebraska: Trans- Syrphidae of Nebraska, with descrip- Ratcliffe, B.C., 1990a, Prairie scarabs- actions of the Nebraska Academy of tions of new species from Nebraska The Nebraska perspective: The Sciences, v. 13, p. 93-98. and Colorado: University Studies, v. American Entomologist, v. 36, p. 28- Smith, H.T.V., 1965, Dune morphology 22, p. 119-162. 35. and chronology in central and west- Williams, D.D., and Miller, D.D., 1952, Ratcliffe, B.C., 1990b, Tiger beetles: ern Nebraska: Journal of Geology, v. A report on Drosophila collections NEBRASKAland Magazine, v. 68, p. 73, p. 557-578. in Nebraska: Bulletin of the Univer- 50-51. Swenk, M.H., 1907, The bees of Ne- sity of Nebraska State Museum, v. 3, Ratcliffe, B.C., 1991 a. The scarab beetles braska—Ill: Entomological News, v. p. 1-19. of Nebraska: Bulletin of the Univer- 18, p. 293-300. Williams, T.A., 1910, The Aphidiidae of sity of Nebraska State Museum, v. Swenk, M.H., and Cockerell, T.D.A., Nebraska: University Studies, v. 10, 12, p. 1-333. 1907a, The bees of Nebraska—I: En- p. 85-175. Ratcliffe, B.C., 1991b, The Passalidae and tomological News, v. 18, p.51-58. Wright, Jr., H.E., 1970, Vegetational his- Lucanidae (Insecta: Coleoptera) of Swenk, M.H., and Cockerell, T.D.A., tory of the Central Plains, in Dort, Nebraska: Great Plains Research, v. 1907b, The bees of Nebraska-11: En- Jr., W., and Jones, Jr., J.K., eds.. Pleis- 1, p. 249-282. tomological News, v. 18, p. 178-187. tocene and Recent Environments of Ratcliffe, B.C., 1991c, Dragonflies: Tate, H.D., and Gates, D.B., 1944, The the Central Great Plains: Lawrence, NEBRASKAland Magazine, v. 69, p. mosquitoes of Nebraska: University University of Kansas Press. 50-51. of Nebraska Agricultural Experiment Zimmer, J.T., 1911, The Pentatomidae of Ratcliffe, B.C., 1992, Swallowtails: Station Research Bulletin No. 133, Nebraska: University Studies, v. 11, NEBRASKAland Magazine, v. 70, p. p. 1-25. p. 219-251. 50-51.

Grasshoppers are widespread in the grasslands of the Sand Hills.