Mole crickets are members of the insect

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Gryllotalpidae, in the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...

Orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grassh ...

(grasshoppers, locusts, and crickets). Mole crickets are cylindrical-bodied,

fossorial A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees. Prehistoric evid ...

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...

s about long as adults, with small

eyes Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and con ...

and shovel-like fore limbs highly developed for burrowing. They are present in many parts of the world and where they have arrived in new regions, may become agricultural pests. Mole crickets have three life stages: eggs,

nymphs A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...

, and adults. Most of their lives in these stages are spent underground, but adults have wings and disperse in the breeding season. They vary in their diet: some species are herbivores, mainly feeding on roots; others are omnivores, including worms and grubs in their diet; and a few are largely predatory. Male mole crickets have an exceptionally loud song; they sing from a burrow that opens out into the air in the shape of an exponential horn. The song is an almost pure tone, modulated into chirps. It is used to attract females, either for mating, or for indicating favourable habitats for them to lay their eggs. In Zambia, mole crickets are thought to bring good fortune, while in Latin America, they are said to predict rain. In Florida, where '' Neoscapteriscus'' mole crickets are not native, they are considered pests, and various biological controls have been used. ''

Gryllotalpa ''Gryllotalpa'' is a genus of insects in the mole cricket family Gryllotalpidae. Species The ''Orthoptera Species File'' lists a number species, including cryptic species groups (indicated with a * and often distinguished only by song pattern ...

'' species have been used as food in West Java, Vietnam, Thailand, Laos, and the Philippines.

Description

Mole crickets vary in size and appearance, but most of them are of moderate size for an insect, typically between long as adults. They are adapted for underground life and are cylindrical in shape and covered with fine, dense hairs. The head, fore limbs, and

prothorax The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum ( ventral), and the propleuron ( lateral) o ...

are heavily

sclerotised Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which qu ...

, but the abdomen is rather soft. The head bears two threadlike antennae and a pair of beady eyes. The two pairs of wings are folded flat over the abdomen; in most species, the fore wings are short and rounded and the hind wings are membranous and reach or exceed the tip of the abdomen; however, in some species, the hind wings are reduced in size and the insect is unable to fly. The fore legs are flattened for digging, but the hind legs are shaped somewhat like the legs of a true cricket; however, these limbs are more adapted for pushing soil, rather than leaping, which they do rarely and poorly. The nymphs resemble the adults apart from the absence of wings and genitalia; the wing pads become larger after each successive moult.

Taxonomy and phylogeny

The Gryllotalpidae are a

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...

group in the order Orthoptera (grasshoppers, locusts, and crickets).

Cladistic analysis Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...

of mole cricket morphology in 2015 identified six tribes, of which four were then new: Indioscaptorini (Scapteriscinae), Triamescaptorini, Gryllotalpellini and Neocurtillini ( Gryllotalpinae), and two existing tribes, Scapteriscini and Gryllotalpini, are revised. The group name is derived straightforwardly from Latin ''gryllus'', cricket, and ''talpa'', mole. Within the extant subfamilies, genera include:

Gryllotalpinae

* tribe Gryllotalpellini Cadena-Castañeda, 2015 (South America) ** '' Gryllotalpella'' * tribe Gryllotalpini Leach, 1815 (World-wide) ** ''

'' * tribe Neocurtillini Cadena-Castañeda, 2015 (Americas) ** '' Leptocurtilla'' ** ''

Neocurtilla ''Neocurtilla'' is a genus of northern mole crickets in the family Gryllotalpidae Mole crickets are members of the insect family Gryllotalpidae, in the order Orthoptera (grasshoppers, locusts, and crickets). Mole crickets are cylindrical-b ...

'' * tribe Triamescaptorini Cadena-Castañeda, 2015 (New Zealand) ** '' Triamescaptor'' * tribe ''incertae sedis'' ** †genus '' Pterotriamescaptor'' ** †genus '' Burmagryllotalpa''

Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...

, Myanmar,

Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...

Scapteriscinae

* tribe Indioscaptorini Cadena-Castañeda, 2015 (Indian Subcontinent) ** '' Indioscaptor'' * tribe Scapteriscini Zeuner, 1939 (Americas) ** ''

Scapteriscus ''Scapteriscus'' is a genus of insects in the family Gryllotalpidae, the mole crickets. Members of the genus are called two-clawed mole crickets.Neoscapteriscus''

†subfamily Marchandiinae

* †'' Archaeogryllotalpoides'' Martins-Neto 1991

Crato Formation The Crato Formation is a geologic formation of Early Cretaceous (Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly during ...

, Brazil,

Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...

* †'' Cratotetraspinus'' Martins-Neto 1997 Crato Formation, Brazil, Aptian * †'' Marchandia'' Perrichot et al. 2002 Charentese amber, France, Cenomanian * †'' Palaeoscapteriscops'' Martins-Neto 1991 Crato Formation, Brazil, Aptian

''Incertae sedis''

* †'' Tresdigitus rectanguli'' Xu, Fang & Wang, 2020 Burmese amber, Myanmar, Cenomanian Mole cricket fossils are rare. A

stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

fossil, ''Cratotetraspinus'', is known from the

Lower Cretaceous Lower may refer to: * Lower (surname) * Lower Township, New Jersey *Lower Receiver (firearms) * Lower Wick Gloucestershire, England See also *Nizhny Nizhny (russian: Ни́жний; masculine), Nizhnyaya (; feminine), or Nizhneye (russian: Н� ...

of Brazil. Two specimens of ''Marchandia magnifica'' in

amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In M ...

have been found in the Lower Cretaceous of Charente-Maritime in France. They are somewhat more abundant in the

Tertiary Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...

amber of the Baltic and Dominican regions; impressions are found in Europe and the American

Green River Formation The Green River Formation is an Eocene geologic formation that records the sedimentation in a group of intermountain lakes in three basins along the present-day Green River in Colorado, Wyoming, and Utah. The sediments are deposited in very f ...

Convergent evolution

Mole crickets are not closely related to the "pygmy mole crickets", the Tridactyloidea, which are in the grasshopper suborder

Caelifera The Caelifera are a suborder of orthopteran insects. They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers (Tetrigoidea) and pygmy mole crickets ( Tridactyloidea). ...

rather than the cricket suborder

Ensifera Ensifera is a suborder of insects that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and Cooloola monsters. This and the suborder Caelifera (grassh ...

. The two groups, and indeed their resemblance in form to the

mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...

ian mole family

Talpidae The family Talpidae () includes the moles (some of whom are called shrew moles and desmans) who are small insectivorous mammals of the order Eulipotyphla. Talpids are all digging animals to various degrees: moles are completely subterranean ...

with their powerful front limbs, form an example of

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...

, both developing adaptations for burrowing.

Behavior

Adults of most species of mole cricket can

fly Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...

powerfully, if not with agility, but males do so infrequently. The females typically take wing soon after sunset, and are attracted to areas where males are calling, which they do for about an hour after sunset. This may be to mate, or they may be influenced by the suitability of the habitat for egg-laying, as demonstrated by the number of males present and calling in the vicinity.

Life cycle

Mole crickets undergo incomplete metamorphosis; when

hatch from eggs, they increasingly resemble the adult form as they grow and pass through a series of up to 10 moults. After mating, a period of 1-2 weeks may occur before the female starts laying eggs. She burrows into the soil to a depth of , ( has been seen in the laboratory), and lays a clutch of 25 to 60 eggs. ''Neoscapteriscus'' females then retire, sealing the entrance passage, but in ''Gryllotalpa'' and ''Neocurtilla'' species, the female has been observed to remain in an adjoining chamber to tend the clutch. Further clutches may follow over several months, according to species. Eggs must be laid in moist ground, and many nymphs die because of insufficient moisture in the soil. The eggs hatch in a few weeks, and as they grow, the nymphs consume a great deal of plant material either underground or on the surface. The adults of some species of mole crickets may move as far as during the breeding season. Mole crickets are active most of the year, but overwinter as nymphs or adults in cooler climates, resuming activity in the spring.

Burrowing

Fossorial front leg of Mole Cricket IMG 5038a

Mole crickets live almost entirely below ground, digging tunnels of different kinds for the major functions of life, including feeding, escape from predators, attracting a mate (by singing), mating, and raising of young. Their main tunnels are used for feeding and for escape; they can dig themselves under ground very rapidly, and can move along existing tunnels at high speed both forwards and backwards. Their digging technique is to force the soil to either side with their powerful, shovel-like fore limbs, which are broad, flattened, toothed, and heavily sclerotised (the

cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...

is hardened and darkened). Males attract mates by constructing specially shaped tunnels in which they sing. Mating takes place in the male's burrow; the male may widen a tunnel to make room for the female to mount, though in some species, mating is tail-to-tail. Females lay their eggs either in their normal burrows or in specially dug brood chambers, which are sealed when complete in the case of the genus ''Neoscapteriscus'' or not sealed in the case of genera ''Gryllotalpa'' and ''Neocurtilla''.

Song

Male mole crickets sing by

stridulating Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...

, always under ground. In '' Gryllotalpa gryllotalpa'', the song is based on an almost pure tone at 3.5 kHz, loud enough to make the ground vibrate 20 cm all round the burrow; in fact, the song is unique in each species. In ''G. gryllotalpa'', the burrow is somewhat roughly sculpted; in '' G. vineae'', the burrow is smooth and carefully shaped, with no irregularities larger than 1 mm. In both species, the burrow has two openings at the soil surface; at the other end is a constriction, then a resonating bulb, and then an escape tunnel. A burrow is used for at least a week. The male positions himself head down with his head in the bulb, and his tail is near the fork in the tunnel. Mole crickets stridulate like other crickets by scraping the rear edge of the left fore wing, which forms a

plectrum A plectrum is a small flat tool used for plucking or strumming of a stringed instrument. For hand-held instruments such as guitars and mandolins, the plectrum is often called a pick and is held as a separate tool in the player's hand. In har ...

, against the lower surface of the right fore wing, which has a ratchet-like series of asymmetric teeth; the more acute edges face backwards, as do those of the plectrum. The plectrum can move forward with little resistance, but moving it backwards makes it catch each tooth, setting up a vibration in both wings. The sound-producing stroke is the raising (levation) of the wings. The resulting song resembles the result of modulating a pure tone with a 66-Hz wave to form regular chirps. In ''G. vineae'', the wing levator muscle, which weighs 50 mg, can deliver 3.5 milliwatts of mechanical power; ''G. gryllotalpa'' can deliver about 1 milliwatt. ''G. vineae'' produces an exceptionally loud song from half an hour after sunset, continuing for an hour; it can be heard up to 600 m away. At a distance of 1 m from the burrow, the sound has a mean power over the stridulation cycle up to 88

decibel The decibel (symbol: dB) is a relative unit of measurement equal to one tenth of a bel (B). It expresses the ratio of two values of a power or root-power quantity on a logarithmic scale. Two signals whose levels differ by one decibel have a ...

s; the loudest recorded peak power was about 92 decibels; at the mouths of the burrow, the sound reaches around 115 decibels. ''G. gryllotalpa'' can deliver a peak sound pressure of 72 decibels and a mean of about 66 decibels. The throat of the horn appears to be tuned (offering low inductive reactance), making the burrow radiate sound efficiently; the efficiency increases when the burrow is wet and absorbs less sound. Mole crickets are the only insects that construct a sound-producing apparatus. Given the known sensitivity of a cricket's hearing (60 decibels), a night-flying ''G. vineae'' female should be able to detect the male's song at a range of 30 m; this compares to about 5 m for a typical ''Gryllus'' cricket that does not construct a burrow. The loudness of the song is correlated with the size of the male and the quality of the habitat, both indicators of male attractiveness. The loudest males may attract 20 females in one evening, while a quieter male may attract none. This behaviour enables acoustic trapping; females can be trapped in large numbers by broadcasting a male's song very loudly.

Food

Mole crickets vary in their diets; some like the tawny mole cricket are

herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...

s, others are

omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nu ...

s, feeding on

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...

worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and no eyes (though not always). Worms vary in size from microscopic to over in length for marine polychaete wor ...

s, roots, and grasses, and others like the southern mole cricket are mainly predacious. They leave their burrows at night to forage for leaves and stems, which they drag underground before consumption, as well as consuming roots underground.

Predators, parasites, and pathogens

Besides

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...

toad Toad is a common name for certain frogs, especially of the family Bufonidae, that are characterized by dry, leathery skin, short legs, and large bumps covering the parotoid glands. A distinction between frogs and toads is not made in scient ...

s, and insectivorous mammals, the predators of mole crickets include subterranean

assassin bugs The Reduviidae are a large cosmopolitan family of the order Hemiptera (true bugs). Among the Hemiptera and together with the Nabidae almost all species are terrestrial ambush predators: most other predatory Hemiptera are aquatic. The main examp ...

wolf spiders Wolf spiders are members of the family Lycosidae (). They are robust and agile hunters with excellent eyesight. They live mostly in solitude, hunt alone, and do not spin webs. Some are opportunistic hunters, pouncing upon prey as they find it or ...

, and various

beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...

s. The South American nematode '' Steinernema scapterisci'' kills ''Neocapteriscus'' mole crickets by introducing

bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...

into their bodies, causing an overwhelming infection. ''Steinernema neocurtillae'' is native to Florida and attacks native ''Neocurtilla hexadactyla'' mole crickets.

Parasitoid wasps Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causi ...

of the genus ''Larra'' (Hymenoptera:

Crabronidae The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over 9 ...

) attack mole crickets, the female laying an egg on the external surface of the mole cricket, and the larva developing externally on the mole cricket host. ''Ormia depleta'' (Diptera:

Tachinidae The Tachinidae are a large and variable family of true fly, flies within the insect order Fly, Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in t ...

) is a specialized parasitoid of mole crickets in the genus '' Neoscapteriscus''; the fly's larvae hatch from eggs inside her abdomen; she is attracted by the call of the male mole cricket and deposits a larva or more on any mole cricket individual (just as many females as males) with which she comes in contact. Specialist predators of mole cricket eggs in China and Japan include the bombardier beetle ''Stenaptinus jessoensis'', whereas in South America, they include the bombardier beetle ''Pheropsophus aequinoctialis'' (Coleoptera:

Carabidae Ground beetles are a large, cosmopolitan family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it is one of the 10 most species-rich animal fami ...

); the adult beetle lays eggs near the burrows of mole crickets, and the beetle larvae find their way to the egg chamber and eat the eggs. Fungal diseases can devastate mole cricket populations during winters with sudden rises of temperature and thaws. The fungus ''

Beauveria bassiana ''Beauveria bassiana'' is a fungus that grows naturally in soils throughout the world and acts as a parasite on various arthropod species, causing white muscardine disease; it thus belongs to the entomopathogenic fungi. It is used as a biologic ...

'' can overwhelm adult mole crickets and several other fungal, microsporidian, and viral pathogens have been identified. Mole crickets evade predators by living below ground, and vigorously burrowing if disturbed at the surface. As a last-ditch defence, they eject a foul-smelling brown liquid from their anal glands when captured; they can also bite.

Distribution

Mole crickets are relatively common, but because they are

nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...

and spend nearly all their lives under ground in extensive tunnel systems, they are rarely seen. They inhabit agricultural fields and grassy areas. They are present in every continent except Antarctica; by 2014, 107 species had been described and more species are likely to be discovered, especially in Asia. '' Neoscapteriscus didactylus'' is a pest species, originating in South America; it has spread to the West Indies and New South Wales in Australia. '' Gryllotalpa africana'' is a major pest in South Africa; other ''

'' species are widely distributed in Europe, Asia, and Australia. They are native to Britain (as to Western Europe), but the former population of ''G. gryllotalpa'' may now be extinct in mainland Britain, surviving in the

Channel Islands The Channel Islands ( nrf, Îles d'la Manche; french: îles Anglo-Normandes or ''îles de la Manche'') are an archipelago in the English Channel, off the French coast of Normandy. They include two Crown Dependencies: the Bailiwick of Jersey, ...

Invasive mole crickets and their biological control

Invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...

are those that cause harm in their newly occupied area, where

biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...

may be attempted. The first-detected invasive mole cricket species was ''Neoscapteriscus didactylus'', a South American species reported as a pest in St. Vincent, West Indies, as early as 1837; by 1900, it was a major agricultural pest in Puerto Rico. It had probably slowly expanded its range northwards, island by island, from South America. The only biological control program against ''N. didactylus'' was in Puerto Rico, and it succeeded in establishing the parasitoid wasp ''

Larra bicolor ''Larra bicolor'' is a parasitoid wasp native to South America. It was introduced into Florida as a biological pest control of invasive mole crickets. Description Adult females of this species are about 22 mm long, with the males somew ...

'' from Amazonian Brazil. In 2001, ''N. didactylus'' in Puerto Rico seemed to be a pest only in irrigated crops and turf. Small-scale experimental applications of the nematode '' Steinernema scapterisci'' were made in irrigated turf, but survival of the nematode was poor. Very much later, this same species was reported as a pest in Queensland, Australia, presumably arriving by ship or plane. The next-detected invasive species was in the late 19th century in Hawaii, probably by ship. It was named as ''Gryllotalpa africana'', but much later as perhaps ''G. orientalis''. It was not identified as ''Gryllotalpa krishnani'' until 2020.Frank J.H. 2020. The identity of the adventive ''Gryllotalpa'' Latreille species (Orthoptera: Gryllotalpidae) in Hawaii, with illustration of male genitalia of ''G. orientalis'' Burmeister. Insecta Mundi 0747: 1-8. It attacked sugarcane and was targeted with ''Larra polita'' from the Philippines in 1925, apparently successfully.Frank, J.H., Leppla, N.C. 2008. Mole crickets (Orthoptera: Gryllotalpidae) and their biological control, pp. 2442-2449. In Capinera, J. L. (ed.). Encyclopedia of Entomology. Springer Science and Business Media. The next detection was in Georgia, USA, and at that time was assumed to be ''N. didactylus'' from the West Indies. It was, in fact, three South American ''Neoscapteriscus'' species, ''N. abbreviatus'', ''N . vicinus'', and ''N. borellii'', probably having arrived in ship ballast. They caused major problems for decades as they spread in the Southeastern USA.''Scapteriscus'' mole cricket populations had built up since the early decades of the 20th century and damaged pastures, lawns, playing fields, and vegetable crops. From the late 1940s,

chlordane Chlordane, or chlordan, is an organochlorine compound that was used as a pesticide. It is a white solid. In the United States, chlordane was used for termite-treatment of approximately 30 million homes until it was banned in 1988. Chlordane was ...

had been the insecticide of choice to control them, but when chlordane was banned by the U.S. EPA in the 1970s, ranchers were left with no economic and effective control method. Especially to aid Florida ranchers, a project that became known as the UF/IFAS Mole Cricket Research Program was initiated in 1978. In 1985, a multiple-authored report was published on accomplishments. In 1988, an account was published on prospects for biological control, and in 1996 an account of promising results with biological control. The program ended in 2004 after 25 years of running monitoring stations, and in 2006 a summary publication announced success: a 95% reduction in mole cricket numbers in northern Florida, with biological control agents spreading potentially to all parts of Florida. Efforts to use ''Larra bicolor'' as a biological control agent in Florida began by importing a stock from Puerto Rico. It became established in a small area of southeastern Florida, but had little effect on ''Neoscapteriscus'' populations. A stock from Bolivia became established in northern Florida and spread widely (with some help) to most of the rest of the state and neighboring states. Its survival depends upon the availability of suitable nectar sources. Once gravid female ''Ormia depleta'' flies were found to be attracted to the song of ''Neoscapteriscus'' males in South America, a path to trap these flies with synthetic mole cricket songs was opened. Experimentation then led to a rearing method. Laborious rearing of over 10,000 flies on mole cricket hosts allowed releases of living fly pupae at many sites in Florida from the far northwest to the far south, mainly on golf courses, and mainly in 1989-1991. Populations were established, began to spread, and were monitored by use of synthetic mole cricket song. Eventually, the flies were found to have a continuous population from about 29°N then south to Miami, but the flies failed to survive the winter north of about 29°N. Shipment and release of the flies to states north of Florida was thus a wasted effort. As the flies had been imported from 23°S in Brazil and could not overwinter north of 29°N, whether flies from 30°S in Brazil might survive better in northern Florida was investigated in 1999, but they did not. The third biological control agent to target ''Neoscapteriscus'' in Florida was the South American nematode ''Steinernema scapterisci''. Small-scale releases proved it could persist for years in mole cricket-infested sandy Florida soils. Its use as a biopesticide against ''Neoscapteriscus'' was patented, making it attractive to industry. Industrial-scale production on artificial diet allowed large-scale trial applications in pastures and on golf courses, which succeeded in establishing populations in several counties, and these populations spread, but sales were disappointing, and the product was withdrawn from the market in 2014. Although experimental application was made in states north of Florida, only in southern Georgia was establishment of the nematode verified, suggesting little interest in the other states.

As pests

Larra bicolor in Paynes Prairie, Florida

The main damage done by mole crickets is as a result of their burrowing activities. As they tunnel through the top few centimetres of soil, they push the ground up in little ridges, increasing evaporation of surface moisture, disturbing germinating seeds, and damaging the delicate young roots of seedlings. They are also injurious to turf and pasture grasses as they feed on their roots, leaving the plants prone to drying out and damage by use. In their native lands, mole crickets have natural enemies that keep them under control. This is not the case when they have been accidentally introduced to other parts of the world. In Florida from the 1940s through the 1980s, they were considered

pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...

and were described as "a serious problem". Their population densities have since declined greatly. A University of Florida entomology report suggests that South American ''Neoscapteriscus'' mole crickets may have entered the United States at

Brunswick, Georgia Brunswick () is a city in and the county seat of Glynn County in the U.S. state of Georgia. As the primary urban and economic center of the lower southeast portion of Georgia, it is the second-largest urban area on the Georgia coastline after S ...

, in ship's ballast from southern South America around 1899, but were at that time mistakenly believed to be from the West Indies. One possible remedy was

biological pest control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...

using the parasitoidal wasps ''Larra bicolor''. Another remedy that has been successfully applied is use of the parasitic nematode '' Steinernema scapterisci''. When this is applied in strips across grassland, it spreads throughout the pasture (and potentially beyond) within a few months and not only controls the mole crickets, but also remains infective in the soil for future years.

In human culture

Fotothek df tg 0007646 Landwirtschaft ^ Pflanzenbau ^ Maulwurf

Folklore

Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most central point. Its neighbours are t ...

, ''Gryllotalpa africana'' is held to bring good fortune to anyone who sees it. In Latin America, ''Scapteriscus'' and ''

'' mole crickets are said to predict rain when they dig into the ground. In Japan in the past, they seem to have been associated with the worms/corpses/bugs that announce a person's sins to heaven in the Koshin/Koushin belief.

As food

''Gryllotalpa'' mole crickets have sometimes been used as food in West Java and Vietnam. In

Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is b ...

, mole crickets ( th, กระชอน) are valued as food in

Isan Northeast Thailand or Isan ( Isan/ th, อีสาน, ; lo, ອີສານ; also written as Isaan, Isarn, Issarn, Issan, Esan, or Esarn; from Pali ''īsānna'' or Sanskrit ईशान्य ''īśānya'' "northeast") consists of 20 prov ...

. They are usually eaten fried along with

sticky rice Glutinous rice (''Oryza sativa var. glutinosa''; also called sticky rice, sweet rice or waxy rice) is a type of rice grown mainly in Southeast and East Asia, and the northeastern regions of South Asia, which has opaque grains, very low amylose ...

.Thai Insect Recipe: Dry Fried Crickets คั่วแมลงกระชอน
/ref> In the Philippines, they are served as a delicacy called ''camaro'' in the province of

Pampanga Pampanga, officially the Province of Pampanga ( pam, Lalawigan ning Pampanga; tl, Lalawigan ng Pampanga ), is a province in the Central Luzon region of the Philippines. Lying on the northern shore of Manila Bay, Pampanga is bordered by Tarlac ...

and are a tourist attraction. They are also served in parts of Northern

Luzon Luzon (; ) is the largest and most populous island in the Philippines. Located in the northern portion of the Philippines archipelago, it is the economic and political center of the nation, being home to the country's capital city, Manila, as ...

Notes

References

External links

Mole Cricket
at Australian Museum *Houston, Terry (2011
Information sheet: Mole Crickets
at Western Australian Museum *Prendergast, Amy (2012)
Solving the Mystery of the Hidden Callers of the Night
" at Australian Wildlife Secrets

at

University of Florida The University of Florida (Florida or UF) is a public land-grant research university in Gainesville, Florida. It is a senior member of the State University System of Florida, traces its origins to 1853, and has operated continuously on its ...

Institute of Food and Agricultural Sciences The University of Florida Institute of Food and Agricultural Sciences (UF/IFAS) is a teaching, research and Extension scientific organization focused on agriculture and natural resources. It is a partnership of federal, state, and county governmen ...

* On the

''Featured Creatures'' website
mole crickets, ''Scapteriscus'' spp.

Bug-a-Boo’s or Grubs Up
{{Taxonbar, from=Q199765 Gryllotalpidae Edible insects Extant Valanginian first appearances