adults fly, so they disperse readily. It eats roots and leaves of grass, especially of Florida’s pasture- and turfgrasses; it destroys tomato, cabbage, eggplant, and bell pepper seedlings. Like the southern mole cricket, it also kills plants by disturbing their roots so that they become desiccated. Like the tawny mole cricket, the shortwinged mole cricket is a damaging herbivore, but it cannot fly and has a restricted distribution. The southern mole cricket has spread widely, but is more of a predator than a phytophage. The specialist native wasp Larra ana- Downloaded from https://academic.oup.com/ae/article-abstract/52/3/138/2474789 by University of Florida user on 11 December 2019 lis F. and the nematode Steinernema neocurtillae, which attack the northern mole cricket, do no harm to the invaders. Generalist natural enemies attack the invaders, but do not inflict enough mortality to control them (Hudson et al. 1988). When these three invaders became problems in Florida in 1899–1924, poisoned baits were used against them. Baits had to be applied year after year, and yet the mole crickets spread. In 1940, a federal emergency relief program gave 1,258 tons of arseni- cal bait to Florida vegetable growers in 12 counties. A low-key biological control research program was J. H. Frank and T. J. Walker conducted for a few years in the 1940s, but was terminated when chlordane was found to control mole crickets cheaply and persistently. In the 1970s, chlordane was banned by the U.S. EPA because of the environmental harm it caused. Florida cattle ranchers were devastated because they could not afford the more expensive chemicals that were available for use against mole crickets in turf and vegetables, and they called on the state legislature to act. The legislature’s response was to earmark part of the funds provided annually to the University of Florida’s Institute of Food and Agricultural Science (UF/IFAS) for mole cricket research. Earmarking was passed down as a man- date to the UF/IFAS Entomology and Nematology Department. In 1978 the UF/IFAS Mole Cricket Research Program was born, and it became the department’s flagship program because a portion of its state funding was mandated to the program. Fig. 1. An adult tawny Fourteen Entomology/Nematology faculty mem- mole cricket, one of the ole crickets are strange insects. Their bers participated: initially C. S. Barfield, D. G. three pests. Photo: Lyle powerful forelegs let them bury them- Buss Boucias, P. G. Koehler, E. L. Matheny, J. L. Nation, Mselves in sand in seconds. Like moles, G. C. Smart, D. E. Short, S. J. Yu, and T. J. Walker they live in the ground. In spring, winged adults in Gainesville; J. A. Reinert in Ft. Lauderdale; and emerge from the ground and fly in swarms after D. J. Schuster in Bradenton; and later R. I. Sailer, sundown, looking for mates and new ground where J. H. Frank, and F. D. Bennett in Gainesville. they can produce offspring. Males tune their bur- Most of these faculty later retired, left Florida, or row entrances as loudspeakers that amplify and dropped out when funding ceased. T. J. Walker direct their calling songs skyward. The louder was coordinator between 1979 and 1985; J. H. they sing, the more females (and competing males) Frank became coordinator in 1985. Postdoctoral they attract. researchers with the program were H. G. Fowler There are four species of mole crickets in (1982–1986), W. G. Hudson (1985–1988), and J. P. Florida. The northern mole cricket, Neocurtilla Parkman (1988–1996). Many student researchers hexadactyla (Perty) (subfamily Gryllotalpinae), is worked under the program. a native species, not a pest, and not closely related The new program investigated mole cricket to the three South American invaders. The invaders origins, life cycles, behavior, physiology, ecology, are the shortwinged mole cricket (Scapteriscus ab- sampling methods, toxicology, and pathology breviatus Scudder), southern mole cricket (S. borel- (Walker 1985). Earlier “basic” research on mole lii Giglio-Tos), and tawny mole cricket (S. vicinus cricket song paid off by making it possible to Scudder; subfamily Scapteriscinae). All are pests, produce synthetic songs that were used to bait but the tawny mole cricket (Fig. 1) is the worst. Its traps to sample flying mole crickets (Walker 138 American Entomologist • Fall 2006 1982). Trapping stations were established at seven Fig. 2. A sound trap points in Florida to mea- for monitoring mole sure long-term population cricket populations, change (Fig. 2). Program and Tom Walker, members realized that its designer. These permanent control was sound traps are in likely to be accomplished pairs (one for tawny only by classical biologi- mole crickets, one for southern mole cal control, but they also crickets, at each investigated chemical con- location). They were Downloaded from https://academic.oup.com/ae/article-abstract/52/3/138/2474789 by University of Florida user on 11 December 2019 trol and relative toler- operated nightly for ance of grass species and 25 years. Photo: Lyle varieties (Walker 1985). Buss “Basic” research on origin of the pests was beneficial in detecting the places in South America where biological control agents the underside of the mole cricket and departs. The might be obtained and imported into Florida mole cricket revives and resumes its usual activities. (Walker and Nickle 1981). Biocontrol agents were The egg hatches within a week, and the wasp larva imported from southern South America in the mid- attaches to the mole cricket and begins feeding. to late 1980s. When almost full-grown, the larva kills its host and In the 1990s, almost all funds for mole cricket consumes the remains. It then makes a cocoon in biocontrol dried up, and the program struggled the ground and pupates. The pupa remains under- to progress. Biological control agents, however, ground, insulated from extreme temperatures, for had been released and a monitoring system was in weeks or, in winter, for months (Castner 1988). place. Under other circumstances, more might have Because 1981–1983 releases had not succeeded been accomplished, but here we describe success in central or northern Florida, R. I. Sailer and F. D. for a part of north-central Florida, knowing that Bennett speculated that the wasps might be climati- the three biological control agents discussed here cally maladapted because they had come from the are spreading success elsewhere. equator. Bennett found better adapted wasps of this same species at cooler sites in Bolivia. Bolivian Larra bicolor wasps were imported and released in 1988/1989 in Larra bicolor F. is a sphecid wasp, native to and near Gainesville. By late 1993, it was evident South America, which attacks only Scapteriscus that the Bolivian strain had become established mole crickets (Menke 1995). Unlike the familiar (Frank et al. 1995). By late 2002, it seemed to hornets and yellow jackets, this wasp is not have spread 135 miles northwest and perhaps as social, does not build or defend nests by stinging far south; it is still spreading, only partially with intruders, and is not a threat to humans. Its larvae assistance (Fig. 4). In time, it is likely to occupy all are parasitoids that feed externally on active mole of Florida. A release at Tifton, GA (>31oN) in 2001 crickets. became established, but a release at Baton Rouge, Although it was known earlier that this wasp LA (<31oN) in 2002 failed. Somehow, the wasp seeks out Scapteriscus mole crickets as food for its arrived in coastal Mississippi (<31oN) (Held 2005). progeny, that knowledge was not put to use until We do not know how far north it may extend its the late 1930s, when Puerto Rican entomologists range; but, because of the wasp’s tropical origin, imported it to their island. Wasps collected on we doubt the range will be much north of 32oN. the equator in Brazil were taken to Puerto Rico This would allow it to inhabit the southernmost and released (Wolcott 1941). The idea was to areas of Georgia, Alabama, Mississippi, Louisiana, establish permanent populations to control pest and southeastern Texas, all of which have severe mole crickets at no further cost. Populations were problems with Scapteriscus mole crickets. established, but nobody measured the effect. In 1981–1983, members of the UF/IFAS program brought wasps from Puerto Rico for release at several places in Florida. These efforts were not very successful. Wasps established a population Fig. 3. Female Larra bicolor only in Ft. Lauderdale and provided little control. attacking a mole They did not spread far afield. cricket. Photo: Behavioral studies on the population of L. Lyle Buss bicolor at Ft. Lauderdale expanded on the Puerto Rican studies. Adult female wasps hunt in daylight for mole cricket adults and large nymphs, entering their burrows and chasing them into the open to attack (Fig. 3). With its sting, the wasp paralyzes its prey for a few minutes. It then lays an egg on American Entomologist • Volume 52, Number 3 139 (Fig. 5). The advantages of this plant were discov- ered in Brazil and Puerto Rico by Wolcott (1941) and seen at Ft. Lauderdale by Castner (1988); the surprise is only that no other plant has yet proven to be more useful. Additional plants are being evaluated. It is most likely that plots of this plant can be manipulated to enhance local populations of the wasp. Details of plot size and spacing are yet to be investigated. Steinernema scapterisci Downloaded from https://academic.oup.com/ae/article-abstract/52/3/138/2474789 by University of Florida user on 11 December 2019 Steinernema scapterisci is a steinernematid nematode native to South America. It lurks in the soil and manages to enter the hemocoel of passing mole crickets, where it releases a specialized bacte- rium that it harbors.