Location-scale mixed models and goodness-of-fit assessment applied to insect ecology.

Survival models have been extensively used to analyse time-until-event data. There is a range of extended models that incorporate different aspects, such as overdispersion/frailty, mixtures, and flexible response functions through semi-parametric models. In this work, we show how a useful tool to assess goodness-of-fit, the half-normal plot of residuals with a simulated envelope, implemented in the hnp package in R, can be used on a location-scale modelling context. We fitted a range of survival models to time-until-event data, where the event was an insect predator attacking a larva in a biological control experiment. We started with the Weibull model and then fitted the exponentiated-Weibull location-scale model with regressors both for the location and scale parameters. We performed variable selection for each model and, by producing half-normal plots with simulated envelopes for the deviance residuals of the model fits, we found that the exponentiated-Weibull fitted the data better. We then included a random effect in the exponentiated-Weibull model to accommodate correlated observations. Finally, we discuss possible implications of the results found in the case study.

Delimiting Strategic Zones for the Development of Fall Armyworm (Lepidoptera: Noctuidae) on Corn in the State of Florida.

The fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), cannot survive prolonged periods of freezing temperatures, thereby limiting where it can overwinter in North America. Climate change is anticipated to reduce the frequency of freeze days in Florida over the decades, with the potential consequence of a significant expansion of the overwintering range, whose northern limit in North America was assessed between 27 and 28°N in the last century. To assess this possibility, the development of the fall armyworm on corn leaves, one of the main host plants in the United States, was determined at five constant temperatures ranging from 14 to 30°C. Based on the development time, the thermal constant and the lower threshold temperature were used to estimate the number of generations of fall armyworm at 42 locations in the state of Florida, from 2006 to 2016. Maps were constructed to provide a visual description of the interpolated data, using GIS (Geographic Information System). The highest number of generations was observed in the counties farther south, an area that showed the highest temperatures during the years and plays a strategic role in maintaining fall armyworm populations in corn fields. Additionally, we conclude that in the absence of freeze periods, the northern limit for fall armyworm overwintering should be between 28 and 29°N.

A Theoretical Approach to Analyze the Parametric Influence on Spatial Patterns of Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) Populations.

Studies of the influence of biological parameters on the spatial distribution of lepidopteran insects can provide useful information for managing agricultural pests, since the larvae of many species cause serious impacts on crops. Computational models to simulate the spatial dynamics of insect populations are increasingly used, because of their efficiency in representing insect movement. In this study, we used a cellular automata model to explore different patterns of population distribution of Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), when the values of two biological parameters that are able to influence the spatial pattern (larval viability and adult longevity) are varied. We mapped the spatial patterns observed as the parameters varied. Additionally, by using population data for S. frugiperda obtained in different hosts under laboratory conditions, we were able to describe the expected spatial patterns occurring in corn, cotton, millet, and soybean crops based on the parameters varied. The results are discussed from the perspective of insect ecology and pest management. We concluded that computational approaches can be important tools to study the relationship between the biological parameters and spatial distributions of lepidopteran insect pests.

Functional Response and Matrix Population Model of Podisus nigrispinus (Dallas, 1851) (Hemiptera: Pentatomidae) fed on Chrysomya putoria (Wiedemann, 1818) (Diptera: Calliphoridae) as Alternative Prey.

Among the predators with high potential for use in biological control, the species of the genus Podisus (Hemiptera: Pentatomidae) have received special attention for laboratory rearing, since they feed on different agricultural and forestry pest insects. However, the type of diet offered to insects in the laboratory may affect the viability of populations, expressed essentially by demographic parameters such as survival and fecundity. This study assessed demographic and development aspects in experimental populations of Podisus nigrispinus (Dallas, 1851) fed on larvae of Chrysomya putoria (Wiedemann, 1818) (Diptera: Calliphoridae) as an alternative prey. The demographic parameters fecundity and survival were investigated in life stages of P. nigrispinus with ecological modeling, by applying the Leslie matrix population model, producing histograms of life stages in successive time steps. The functional response of P. nigrispinus was also investigated on seven densities of C. putoria third-instar larvae at 24 and 48 h. The survival of predators that reached adulthood was 65% and the development time from egg to adult was 23.15 days. The predator showed a type III functional response for consumption of C. putoria at 24 and 48 h. The Leslie-matrix simulation of the age structure provided perpetuation of the predator population over time steps and the prey proved to be feasible for use in rearing and maintenance of P. nigrispinus in the laboratory.

Co-Infestation and Spatial Distribution of Bactrocera carambolae and Anastrepha spp. (Diptera: Tephritidae) in Common Guava in the Eastern Amazon.

Field infestation and spatial distribution of introduced Bactrocera carambolae Drew and Hancock and native species of Anastrepha in common guavas [Psidium guajava (L.)] were investigated in the eastern Amazon. Fruit sampling was carried out in the municipalities of Calçoene and Oiapoque in the state of Amapá, Brazil. The frequency distribution of larvae in fruit was fitted to the negative binomial distribution. Anastrepha striata was more abundant in both sampled areas in comparison to Anastrepha fraterculus (Wiedemann) and B. carambolae The frequency distribution analysis of adults revealed an aggregated pattern for B. carambolae as well as for A. fraterculus and Anastrepha striata Schiner, described by the negative binomial distribution. Although the populations of Anastrepha spp. may have suffered some impact due to the presence of B. carambolae, the results are still not robust enough to indicate effective reduction in the abundance of Anastrepha spp. caused by B. carambolae in a general sense. The high degree of aggregation observed for both species suggests interspecific co-occurrence with the simultaneous presence of both species in the analysed fruit. Moreover, a significant fraction of uninfested guavas also indicated absence of competitive displacement.

Induced Cannibalism in Experimental Populations of the Forensic Indicator Chrysomya putoria Wiedemann (Diptera: Calliphoridae).

We analyzed the behavior of third-instars of Chrysomya putoria as potential cannibals in experimental populations. Cannibalism rates were evaluated in three settings observed for 3, 6, 9, and 24 h, placing injured and uninjured larvae of C. putoria together. Our data heavily support that C. putoria larvae behave as cannibals when induced by a wound in another larva, and also after starving for 24 h. The probability of cannibalism increased as a function of time, both in no-choice and in choice experiments evidencing that time is a determining factor for cannibalism induction in C. putoria. However, the treatment combining injured with uninjured larvae showed the highest probability of cannibalism. These results suggest that C. putoria larvae may cannibalize under scarcity of food over long time or the presence of injured larvae. This study is useful to understand the behavior of C. putoria feeding on ephemeral substrates such as carrion or corpses and brings relevant and significant contribution to population ecology of blowflies and also forensic entomology.

Imidacloprid affects the functional response of predator Podisus nigrispinus (Dallas) (Heteroptera: Pentatomidae) to strains of Spodoptera frugiperda (J.E. Smith) on Bt cotton.

Podisus nigrispinus (Dallas) (Heteroptera: Pentatomidae) is one of the most common asopine species in the neotropical region and its occurrence was reported in several countries of South and Central America, as an important biological control agent for many crops. This study was carried out to identify the imidacloprid impacts on the functional response of predator P. nigrispinus fed on Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) strain resistant to lambda-cyhalothrin, on Bt cotton expressing Cry1Ac (Bollgard(®)). Spodoptera frugiperda larvae were used in the following conditions: resistant (1) and susceptible (2) strains to lambda-cyhalothrin fed on Bollgard(®) cotton leaves (DP 404 BG); and resistant (3) and susceptible (4) strains to lambda-cyhalothrin fed on non-genetically modified cotton leaves (cultivar DP4049). The predatory behavior of P. nigrispinus was affected by imidacloprid and the type II asymptotic curve was the one that best described the functional response data. Handling time (T h ) of predator females did not differ among treatments in the presence of imidacloprid. The attack rate did decrease, however, due to an increase in the density of larvae offered. Regardless of the treatment (S. frugiperda strain or cotton cultivar), the predation of P. nigrispinus females on S. frugiperda larvae was significantly lower when exposed to imidacloprid, especially at a density of 16 larvae/predator. The predation behavior of P. nigrispinus on S. frugiperda larvae is affected by the insecticide imidacloprid showing that its applications should be used in cotton crop with caution.

Population dynamics, life stage and ecological modeling in Chrysomya albiceps (Wiedemann) (Diptera: Calliphoridae).

In this study we investigated the population dynamics of Chrysomya albiceps (Wiedemann) with laboratory experiments, employing survival analysis and stage structure mathematical models, emphasizing survival among life stages. The study also assessed the theoretical influence of density dependence and cannibalism during immature stages, on the population dynamics of the species. The survival curves were similar, indicating that populations of C. albiceps exhibit the same pattern of survival among life stages. A strong nonlinear trend was observed, suggesting density dependence, acting during the first life stages of C. albiceps. The time-series simulations produced chaotic oscillations for all life stages, and the cannibalism did not produce qualitative changes in the dynamic behavior. The bifurcation analysis shows that for low values for survival, the population reaches a stable equilibrium, but the cannibalism results in chaotic oscillations practically over all the parametric space. The implications of the patterns of dynamic behavior observed are discussed.

Breeding of Ornidia obesa (Diptera: Syrphidae: Eristalinae) on pig carcasses in Brazil.

Ornidia obesa F. (Diptera: Syrphidae) is usually neglected in forensic entomology, although adults are rather frequent on vertebrate carrion. In this study, conducted in southeastern Brazil in 2008, we used two pig carcasses, one killed by cocaine overdose and the other by shooting, to evaluate mainly the possible influences of the type of death on the larval development of O. obesa in the pig remains. We recorded the breeding of 218 adult specimens of this syrphid fly from the carcass killed by shooting, and none from the carcass killed by cocaine. These observations may open a new perspective for the use of O. obesa in forensic studies, considering its breeding preferences and its complete development on vertebrate carrion.

Fecundity, body size and population dynamics of Chrysomya albiceps (Wiedemann, 1819) (Diptera: Calliphoridae).

In this study, the seasonal variation of fecundity, wing and tibia length were investigated in natural populations of Chrysomya albiceps (Wiedemann, 1819) in an attempt to determine the changes in life history of the species as a function of seasonality. A relative constant temporal trajectory was found for fecundity, wing and tibia length over twenty-four months. Positive correlations between fecundity and wing size, fecundity and tibia size and wing and tibia sizes were observed. The implications of these results for population dynamics of C. albiceps are discussed.

Fecundity, body size and population dynamics of Chrysomya albiceps (Wiedemann, 1819) (Diptera: Calliphoridae)

In this study, the seasonal variation of fecundity, wing and tibia length were investigated in natural populations of Chrysomya albiceps (Wiedemann, 1819) in an attempt to determine the changes in life history of the species as a function of seasonality. A relative constant temporal trajectory was found for fecundity, wing and tibia length over twenty-four months. Positive correlations between fecundity and wing size, fecundity and tibia size and wing and tibia sizes were observed. The implications of these results for population dynamics of C. albiceps are discussed.

Neste estudo, a variação sazonal da fecundidade, comprimentos de asa e tíbia foram investigadas em populações naturais de Chrysomya albiceps (Wiedemann, 1819) como uma forma de determinar as variações bionômicas da espécie em função da sazonalidade. Uma trajetória temporal constante foi encontrada para fecundidade, comprimentos de asa e tíbia durante vinte e quatro meses. Correlações positivas entre fecundidade e comprimento de asa, fecundidade e comprimento de tíbia e comprimentos de asa e tíbia foram observadas. As implicações destes resultados para a dinâmica populacional de C. albiceps foram discutidas.

Sex ratio and dynamic behavior in populations of the exotic blowfly Chrysomya albiceps (Diptera, Calliphoridae).

Sex ratio is an essential component of life history to be considered in population growth. Chrysomya albiceps is a blowfly species with a naturally biased sex ratio. In this study, we evaluated the impact of changes in sex ratio on the dynamic behavior of C. albiceps using a density-dependent mathematical model that incorporated demographic parameters such as survival and fecundity. These parameters were obtained by exponential regression, with survival and fecundity being estimated experimentally as a function of larval density. Bifurcation diagram of the results indicated the evolution of stable equilibrium points as a function of sex ratio. A continually increasing sex ratio yielded a hierarchy of bifurcating stable equilibrium points that evolved into a chaotic regime. The demographic parameters obtained by exponential regression were also changed to maximum and minimum values in order to analyze their influence on dynamic behavior with sex ratio being considered as an independent variable. Bifurcations with periodicity windows between chaos regimes were also found.

Sex ratio and dynamic behavior in populations of the exotic blowfly Chrysomya albiceps (Diptera, Calliphoridae)

Sex ratio is an essential component of life history to be considered in population growth. Chrysomya albiceps is a blowfly species with a naturally biased sex ratio. In this study, we evaluated the impact of changes in sex ratio on the dynamic behavior of C. albiceps using a density-dependent mathematical model that incorporated demographic parameters such as survival and fecundity. These parameters were obtained by exponential regression, with survival and fecundity being estimated experimentally as a function of larval density. Bifurcation diagram of the results indicated the evolution of stable equilibrium points as a function of sex ratio. A continually increasing sex ratio yielded a hierarchy of bifurcating stable equilibrium points that evolved into a chaotic regime. The demographic parameters obtained by exponential regression were also changed to maximum and minimum values in order to analyze their influence on dynamic behavior with sex ratio being considered as an independent variable. Bifurcations with periodicity windows between chaos regimes were also found.

A razão sexual é um componente essencial da biologia de organismos a ser considerado em crescimento populacional. Chrysomya albiceps é uma espécie de mosca varejeira que exibe um desvio na razão sexual em relação à proporção 1:1. Neste estudo avaliamos o impacto de alterações na razão sexual sobre o comportamento dinâmico de C. albiceps utilizando um modelo matemático dependente da densidade que incorpora parâmetros demográficos como sobrevivência e fecundidade. Os parâmetros foram obtidos por regressão exponencial, com sobrevivência e fecundidade sendo estimadas experimentalmente em função da densidade larval. O diagrama de bifurcação gerado pelos resultados indicou a evolução de pontos de equilíbrio estável em função da razão sexual. A sucessão contínua dos valores da razão sexual resultou em uma hierarquia de pontos de equilíbrio estável produzida por bifurcações, resultando em regime caótico. Os parâmetros demográficos obtidos por regressão exponencial foram também mudados para valores máximos e mínimos, a fim de analisar a influência deles sobre o comportamento dinâmico da espécie, tendo a razão sexual como variável independente. Bifurcações com janelas de periodicidade intercaladas com o regime caótico também foram encontradas.

Larval density, temperature and biological aspects of Chrysomya megacephala (Diptera: Calliphoridae) / Densidade larval, temperatura e aspectos biológicos de Chrysomya megacephala (Diptera: Calliphoridae)

The effect of larval density on the survival, fecundity and body size at two temperatures in experimental populations of C. megacephala was studied. No effect from simultaneous influence of density and temperature on life history characteristics of C. megacephala was found. Significant effects of density and temperature on survival, fecundity and body size were observed. The importance of these results for the population dynamics of C. megacephala is discussed.

Analisou-se o efeito de duas densidades larvais e duas temperaturas sobre a sobrevivência, fecundidade e tamanho corpóreo de C. megacephala em condições experimentais. Nenhum efeito simultâneo da densidade e temperatura foi encontrado sobre as variáveis investigadas em C. megacephala. Entretanto, foram observados efeitos isolados significativos da densidade e da temperatura sobre a fecundidade e tamanho corpóreo. A importância desses resultados para a dinâmica populacional de C. megacephala foi discutida.

Dynamics of extinction in coupled populations of the flour beetle Tribolium castaneum.

In this study we analyzed the effect of migration on the persistence time of coupled local populations of Tribolium in different environments. Four treatments were set up to compare different levels of environmental heterogeneity. We established high, low, moderate, and no heterogeneity. These levels were estimated by the different amounts of food offered to each population. To investigate how risk spreading works, a stochastic model for two subpopulations was employed. The high heterogeneity treatment resulted in the longest persistence, even though survival analysis revealed no significant difference among treatments. The magnitude of differences in growth rates among subpopulations is probably associated with persistence.

Dynamics of extinction in coupled populations of the flour beetle Tribolium castaneum

Neste estudo analisamos o efeito da migração sobre o tempo de persistência de populações acopladas de Tribolium em diferentes ambientes. Quatro tratamentos foram estabelecidos para comparar diferentes níveis de heterogeneidade ambiental, alto, moderado, baixo e nulo. Os níveis de heterogeneidade foram estabelecidos por meio de diferentes quantidades de alimento oferecidas a cada população. Para investigar como funciona a expansão de risco entre populações conectadas, um modelo estocástico para duas populações acopladas foi empregado. O tratamento estabelecido para analisar a alta heterogeneidade ambiental foi o que exibiu maior tempo de persistência, apesar da análise de sobrevivência não revelar diferença significativa entre os tratamentos. A magnitude da diferença nas taxas de crescimento entre as populações provavelmente está associada ao tempo de persistência populacional.

Relaçäo entre desempenho e infecçöes por nematódeos gastrintestinais em bovinos Nelore em crescimento / Relationship between performance and infections by gastrointestinal nematodes in growing Nelore cattle

Este estudo teve por objetivo verificar a associaçäo entre o desempenho e a carga parasitária de 28 machos inteiros da raça Nelore (a partir de um ano de idade), naturalmente infectados por nematódeos gastrintestinais. De novembro de 1999 a junho de 2000, os animais foram pesados quinzenalmente, período no qual foram colhidas amostras de fezes e de sangue para a realizaçäo, respectivamente, de exames coprológicos e sangüíneos. Os dados da contagem de ovos por grama de fezes (OPG) ajustaram-se ao modelo de distribuiçäo binomial negativa, indicando que a distribuiçäo de nematódeos no rebanho é agregada. Näo ficou evidenciada associaçäo entre ganho de peso e contagem de OPG, e entre contagem de OPG e volume globular (VG). De 10 coeficientes de correlaçäo entre ganho de peso e VG, oito foram negativos, mas apenas dois significativos (P<0,05). As estimativas de repetibilidade da contagem de OPG, LOG (OPG+1) e VG foram de 0,26, 0,25 e 0,33, respectivamente. Cooperia punctata foi a espécie mais freqüentemente encontrada parasitando os animais. Além dessa espécie, foram detectados os seguintes nematódeos: Haemonchus placei, Haemonchus similis, Trichostrongylus axei, Bunostomum phlebotomum e Oesophagostomum radiatum. O parasitismo por nematódeos gastrintestinais aparentemente näo prejudicou o desenvolvimento dos animais estudados

Competiçäo larval em Chrysomya megacephala (F.) (Diptera: Calliphoridae): efeitos de diferentes níveis de agregaçäo larval sobre estimativas de peso, fecundidade e investimento reprodutivo / Larval competition in Chrysomya megacephala (F.) (Diptera: Calliphoridae): effects of different levels of larval aggregation on estimates of weight, fecundity and reproductive investment

Em insetos que se utilizam de substratos discretos e efêmeros para a alimentação e postura de ovos, o resultados da competição larval por alimento depende da quantidade de recursos e da distribuição espacial dos imaturos nos substratos alimentares. No presente estudo, os resultados da competição larval por alimento em Chrysomya megacephala, em caracteres como peso de fêmeas, fecundidade e investimento reprodutivo, foram diferentes em situações em que o nível de agregação larval (proporção de competidores por qualtidade de alimento) é o mesmo, mas com densidades de competidores e quantidades de alimento proporcionalmente diferentes. Esses resultados indicam que a competição larval pode depender tanto da densidade larval como da quantidade de alimento, em situações diferentes com a mesma proporção de larvas por grama de alimento.

Diffusion model applied to postfeeding larval dispersal blowflies (Diptera: Calliphoridae)

This paper presents a diffusion model of larval dispersal especifically designed to account for particular aspects of postfeeding larval dispersal from the food source in organisms such as blowflies. In these organisms the dispersal of immatures includes two groups of individuals, those that are actively migrating and those that initiated the pupation process. The classical diffusion equation in one dimension was modified to incorporate a function which describes the burying of larvae to become pupae. The analytical solution of this equation predicts oscillatory and monotonic dispersal behaviors, which are observed in experimental populations of blowfly species.

Dynamics of experimental populations of native and introduced blowflies (Diptera: Calliphoridae): mathematical modelling and the transition from asymptotic equilibrium to bounded oscillations

The equilibrium dynamics of native and introduced blowflies is modelled using a density-dependent model of population growth that takes into account important features of the life-history in these flies. A theoretical analysis indicates that the product of maximum fecundity and survival is the primary determinant of the dynamics. Cochliomyia macellaria, a blowfly native to the Americas and the introduced Chrysomya megacephala and Chrysomya putoria, differ in their dynamics in that the first species shows a damping oscillatory behavior leading to a one-point equilibrium, whereas in the last two species population numbers show a two-point limit cycle. Simulations showed that variation in fecundity has a marked effect on the dynamics and indicates the possibility of transitions from one-point equilibrium to bounded oscillations and aperiodic behavior. Variation in survival has much less influence on the dynamics.