Monte Lloyd's research while affiliated with Tri State University and other places
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Publications (15)
An instance has been documented where two 13-year broods of periodical cicadas, XIX and XXIII, occur together in the same woods. Unusually good historical records suggest that this situation arose as the aftermath of hybridization in 1868 between one of the 13-year broods (XIX) and 17-year Brood X, after which the 17-year brood disappeared from the...
Periodical cicada nymphs (Homoptera:Cicadidae:Magicicada spp.) that were collected as they emerged from the ground, isolated, and allowed to eclose in individual containers sometimes developed infective conidia of the fungus Massospora cicadina Peck, but never resting spores. The time required for the cicada's abdominal sclerites to fall off (i.e.,...
Flight capabilities of healthy and fungus infected Magicicada septendecim L. (Homoptera: Cicadidae) were compared using 3 complementary techniques: 1) observations of spontaneous flights, 2) field-tested flights, and 3) tethered flights in which endurance was measured. Spontaneous flight distances are much lower than those obtained on field tested...
Infected periodical cicadas, Magicicada cassini (Cicadidae: Homoptera), that are developing resting spores of the fungus Massospora cicadina (Entomophthorales) invade new habitat (nearby young trees) about as readily as do uninfected cicadas, although this may not be true for another periodical cicada species, M. septendecim. Proportions of the two...
An unusual occurrence of periodical cicadas in a white pine-hemlock plantation in Ohio is documented. Historical information suggests that cicadas invaded this plantation when intermixed deciduous species were available for oviposition. Subsequent studies show that hatching success is far better in deciduous plants than in pines. The reduction in p...
Densities of two allochronic, geographically overlapping populations of periodical cicadas were censused in the eggnest stage in order to test the hypothesis that larger populations can be supported when the root-feeding nymphs are of different ages. In a frequently burned chest-high stand of scrub oak Quercus ilicifolia on Long Island, Magicicada...
Eggs of periodical cicadas (Magicicada spp.) will often die within the twigs where they are laid if the twig withers from excessive damage (called "flagging"). Many of these dead eggs also become stained. Artificially cutting the twigs during embryogenesis results in less egg mortality but more staining than seen in naturally flagged twigs. An expe...
We propose a model, focused on competition among periodical cicadas, which partitions the conventional competition coefficients into three components: how the nymphs affect each other, precisely where they are located, and the disparity between fecundities. The second component depends on having measured the microspatial distribution and determinin...
We postulate that the potential exists in 17-year cicadas to (1) accelerate by four years or (2) decelerate by one year, and that both are developmental aberrations induced by crowding and/or shortage of food owing to extreme competition. Further, we postulate that process (1) can gradually lead to the formation of a new brood, but that process (2)...
A complete count of exuviae from a yard of @?0.1 ha revealed 158,054 emerging periodical cicadas (Magicicada cassini) of which 31% died from faulty eclosion. Mutual interference and/or delay owing to a shortage of vertical perching places appears to be responsible, rather than poor nutrition. We argue that cicadas feeding underground are protected...
The geographical relationships and relative sizes of 17-year periodical cicada broods, along with the fact that 13-year cicadas exist, suggest that 17-year cicadas sometimes undergo a 4-year acceleration and emerge after 13 years (Lloyd and Dybas, 1966). The early growth of 17-year nymphs is strikingly inhibited by 4 years, relative to that of 13-y...
Early growth of 17-year cicadas is much slower than that of the 13-year race, even when the 17-year nymphs are in a warmer climate. It is postulated that 17-year cicadas have a physiological mechanism inhibiting the growth of the early instars, possibly by restricting the season of feeding. Nymphs of the same age in the same forest or orchard are s...
Citations
... Characters such as body size, longevity, and locomotion during feeding are well known to be directly affected by the nutrition available to larvae (Robertson 1963;Chippendale et al. 1993;Partridge & Fowler 1993;Sokolowski et al. 1997;Foley & Luckinbill 2001). It is well known that in many species the environmental conditions of immatures-larvae, nymphs-have permanent effects which persist into the adult stage of life history (Marlatt 1907;Uhlenhuth 1919;Citus 1935;Ryan 1941;Bates 1949;Istock 1966;Southwood 1966;Azam & Anderson 1969;Engelmann 1970;Brown & Chippendale 1973;Wilbur & Collins 1973;Otto 1974;Lloyd & White 1976;Wilbur 1977a;Howard 1978;Pritchard 1978;Steinwascher 1978;Sweeney & Vannote 1978;Vannote 1978;Anderson & Cummins 1979;Dunlap-Pianka et al. 1979;Ward & Cummins 1979;Caligari 1980;Harshman 1982;Marks 1982;Prout & McChesney1985;Semlitsch 1987). Relevant to population dynamics, the adult fertility component of growth can be affected by limiting resources of immatures. ...
... Density information is essential for distinguishing ordinary emergences from possible 'straggler' or off-cycle emergences. Periodical cicadas depend on high population density and predator-swamping (White and Lloyd 1979;Lloyd and White 1980;Karban 1982a,b;Williams et al. 1993;Williams and Simon 1995;Marshall 2001), so records of single or scattered individuals are unlikely to represent sustaining populations. Lacking GPS technology in 2000, we marked presence and absence records on 1:150,000 paper maps (DeLorme 2012), placing our marks relative to the detailed information on road orientation, intersections, and other printed landmarks offered in this map series. ...
... The genetic basis of life cycle length has not been studied because the long life cycles complicate genetic crosses. An early explanation for life cycle control in periodical cicadas proposed a one-locus, two-allele system in which either the 13-or the 17-year cycle is dominant 10,11 . Differences between the two life cycle lengths may be attributable to differences in juvenile developmental rate 12,13 , which may be regulated by one locus or a small number of loci. ...
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... Cicadas cluster in high density in certain places, or are patchy (Simon et al. 1981). Dybas and Lloyd (1974) remarked that, contrary to published concerns from 19 th century cicada researchers, the cicadas adapt surprisingly well to the human disturbance of tree-cutting (see also Collinge [2010] on the effects of habitat fragmentation). ...
... For example, we implemented relatively simple functional forms for competition; however, there is currently a limited understanding of the mechanistic underpinnings of competition. Furthermore, individual periodical cicadas occasionally emerge in years different from the majority of their cohort and instead delay or accelerate their year of emergence [23, 19, 25, 10, 22]. Our forthcoming research implements alternative functional forms for competition as well as exploring this " leakage " phenomenon to determine whether our results apply to models with more general models of cicadas. ...
... 2). Maier (1985) noted that many scattered eastern reports of Brood VI were likely attributable to misidentification or off-cycle emergences as hypothesized by Lloyd and Dybas (1966) and Lloyd and White (1976). Marshall (2001) extended this observation to scattered reports across the general range of periodical cicadas. ...
... Cicada egg developmentvaries among species; for example, in the periodical cicada Magicicada sp., eggs are laid on living branches and develop for 6-8 weeks before hatching, which occurs around early August (White and Lloyd, 1981). In our study, P. plagifera egg's incubation period of 48 days is almost similar to 45-50 days reported by Moalla et al. (1992). ...
... These observations motivated numerous studies in theoretical population biology to understand the reasons why large prime number periods have been selected by evolution, but far fewer studies explaining how the two levels of synchrony are achieved. For prime number selection, the hypothesis [7,8] that limited environmental carrying capacity and predation pressure are responsible was first captured in a mathematical model by Hoppensteadt and Keller [9]. Later models elucidated mechanisms by which single broods occupy disjoint areas [10][11][12]. ...
... Although he did not develop a life-cycle control hypothesis with a 4-year sub-cycle, this could be a key component of the year-counting mechanism. For the genetic basis of two life cycles, Lloyd et al. (1983) and Cox and Carlton (1991) proposed that the 4-year difference is caused by a singlelocus two-allele system dominated by either a 17-year allele (Lloyd et al., 1983) or a 13-year allele (Cox & Carlton, 1991). However, circumstantial evidence that these hypotheses have relied on has been falsified (Marshall, 2001). ...
... For example, Cory and Knight (1937) found an approximately 31% failure rate during heavy emergences in Brood X cicadas from 1936. White et al. (1979) found that up to 31% of the cicadas emerging in extreme numbers under isolated sugar maples in a suburban lawn during the 1976 emergence of Brood XXIII died from failed ecdysis. These observations suggest that in some circumstances, high densities may be a liability. ...
