Larval Distribution and Behavior of Chrysomya rufifacies (Macquart) (Diptera: Calliphoridae) Relative to Other Species on Florida Black Bear (Carnivora: Ursidae) Decomposing Carcasses

Larval interactions of dipteran species, blow flies in particular, were observed and documented daily over time and location on five black bear carcasses in Gainesville, FL, USA, from June 2002 – September 2004. Cochliomyia macellaria (Fabricius) or Chrysomya megacephala (Fabricius) larvae were collected first, after which Chrysomya rufifacies (Macquart) oviposited on the carcasses in multiple locations (i.e., neck, anus, and exposed flesh) not inhabited already by the other blow fly larvae. Within the first week of decomposition, C. rufifacies larvae grew to ≥12 mm, filling the carcasses with thousands of larvae and replacing the other calliphorid larvae either through successful food source competition or by predation. As a result, C. macellaria and C. megacephala were not collected past their third instar feeding stage. The blow fly species, C. megacephala, C. macellaria, Lucilia caeruleiviridis (Macquart), Phormia regina (Meigen), Lucilia sericata (Meigen), and C. rufifacies, completed two developmental cycles in the 88.5-kg carcass. This phenomenon might serve to complicate or prevent the calculation of an accurate postmortem interval.     

The distribution of adult blow-flies (Diptera: Calliphoridae) along an altitudinal gradient in Central Spain

The distribution of Calliphoridae along an altitudinal gradient was investigated in Central Spain using carrion-baited traps. Significant differences were found between elevation and mean abundances of almost all species of blow-flies. Several species of flies could be grouped according to their altitudinal preferences so that samples at high elevations are defined by Calliphora vomitoria and Calliphora vicina while samples at low elevations are defined by two thermophilous species: Lucilia sericata and Chrysomya albiceps. The remaining species show preferences for mid-elevations where wooded areas are more characteristic along the altitudinal gradient. Calliphora vomitoria and Chrysomya albiceps are the most abundant species representing the 87.74 % of all captures. Both species are spatially segregated along the altitudinal gradient. The changing patterns of abundance are discussed in relation to differences in climate conditions along the altitudinal gradient concluding that the environmental variables that influence the seasonality of many species also play an important role to explain the spatial distribution.     

Successional patterns of the insect fauna on a pig carcass in southern Italy and the role of Crematogaster scutellaris (Hymenoptera,Formicidae) as a carrion invader

The sarcosaprophagous fauna plays a key role in organic matter decomposition. Moreover, the biological, ecological and behavioral specificities of the taxa are useful to reconstruct the decay history of a corpse or carcass, often back to the lethal event. Here we report the seasonal succession of the insect fauna on a pig carcass exposed in a rural area in Calabria (southern Italy) during summer 2007 and 2008. The aim is to identify and qualitatively assess the major taxa of forensic importance in this region. The principal fly invaders were Lucilia caesar (L.), L. sericata (Meigen, 1826), Chrysomya albiceps (Wiedemann, 1819), Sarcophaga (Meigen, 1826) spp. and Amobia (Robineau‐Desvoidy, 1830) spp., Musca domestica (L.) and Muscina stabulans (Fallen, 1817). The primary beetle species collected in summer belonged to Dermestidae, Dermestes maculatus (De Geer, 1774) and Cleridae, Necrobia rufipes (De Geer, 1775). This paper also examined the ecological role of ants in the insect succession and describes the evidence of skin injuries directly inflicted by the acrobat ant Crematogaster (Acrocoelia) scutellaris (Olivier, 1791) (Hymenoptera Formicidae) while feeding on pig carrion. Ants belonging to two other species were also collected: Camponotus aethiops (Latreille, 1798) and Tetramorium semilaeve (André, 1881). Ants can invade carcasses and corpses directly, disrupting blowfly egg laying or preying on their larvae. Our data on the carrion faunal composition and role of ants as invaders should be useful for further forensic cases in Calabria (southern Italy). This is among the few reports of ants as forensically relevant species.     

Succession of Carrion Fauna in the Arid Region of San Juan Province, Argentina and Its Forensic Relevance

The succession of carrion fauna and the decomposition stages were studied in the arid environment of San Juan Province, Argentina (31°32??34.7?? S; 68°34??39.4?? W). Two pig carcasses (Sus scrofa) were placed in wire mesh cages, 100?m apart from each other. Each carcass was surrounded by pitfall traps, and a modified Malaise trap was placed above. Daily samplings were carried out to collect the insects present in the carcasses and the traps, and body and environmental temperature were measured. The main colonizer species was Chrysomya albiceps (Wiedemann) and its larvae were dominant over other Diptera. The first adult blowflies emerged at 8.9?days. The time to reach the remains stage was 8?days shorter than for other South American sites. We recorded the early arrival of adult Dermestes maculates De Geer and Dermestes ater De Geer 2?days after the beginning of the assay, and larvae of these species were recorded 4?days after. We determined a 1.5-day error in the postmortem interval estimation using the temperatures measured in the assay and those recorded by the nearest meteorological station.     

Forensic insects attracted to human cadavers in a vehicular environment in Riyadh,Saudi Arabia

Insect fauna attracted to cadavers at the crime scene can be identified and used to estimate the post-mortem interval (PMI). In the current study, insects associated with two human cadavers in a vehicular environment were collected and analysed. The first cadaver was found five days’ post mortem in a garage. The second cadaver was found in a car ten days after his death. The weather conditions were obtained from the nearest weather station located to the scenes of the death. During the study, six adults, 32 larvae and egg batches were collected from case 1 and identified as Chrysomya albiceps Wiedemann (Diptera: Calliphoridae). From the second case, two larvae of Megaselia scalaris Loew (Diptera: Foridae) and seven larvae of Musca domestica L. (Diptera: Muscidae) were identified. mPMI ranged from five to twelve days. This information expands the knowledge on the insect fauna in the vehicular environment, which could be used to assist estimation of the PMI.     

Effect of Lunar Phases,Tides, and Wind Speed on the Abundance of Diptera Calliphoridae in a Mangrove Swamp

Abiotic factors, such as lunar phases and tides, have a significant effect on insect development. Reproduction and immature development are usually interlinked to these abiotic factors. The tide is at its highest levels at full moon or new moon, hindering the feeding of the immature or causing their drowning. The oviposition by adult females is also compromised on these days because much of the available food is submerged. Another important abiotic factor is the wind, which displaces odoriferous particles in the air. Wind speed and direction are important elements to indicate potential sources of food for insects. I report on the effects of lunar phases, tides, and wind speed on the Calliphoridae fauna in mangrove swamps. The different species collected were identified, and the predominant species in the area were quantified. A total of 1,710 flies were collected over a 1-year period. Six Calliphoridae flies, Chloroprocta idioidea (Robineau-Desvoidy), Chrysomya megacephala (Fabricius), Chrysomya albiceps (Wiedemann), Chrysomya putoria (Wiedemann), Cochliomyia macellaria (Fabricius), and Lucilia eximia (Wiedemann) were collected. Data indicated that lunar phases have a significant effect on the abundance of C. albiceps (r?=?0.39, p?<?0.01), and that the variation of the tides also affected the abundance of C. putoria (r?=?0.40, p?<?0.00), C. macellaria (r?=?0.41, p?<?0.00), and C. idioidea (r?=?0.31, p?<?0.04). The wind speed, however, did not affect these species.     

Dispersal and Predation Behavior in Larvae of <Emphasis Type="Italic">Chrysomya albiceps</Emphasis> and <Emphasis Type="Italic">Chrysomya megacephala</Emphasis> (Diptera: Calliphoridae)

Chrysomya albiceps and Chrysomya megacephala are exotic blowfly species known by producing myiasis in humans and other animals and by transmitting pathogens mechanically. C. albiceps stand out by being a facultative predator of other dipteran larvae. In this paper we investigated the influence of larval predation on the dispersal of larvae of C. albiceps and C. megacephala single and double species for three photophases. An experimental acrylic channel graduated and covered with wood shavings was used to observe the larval dispersal. The results showed that C. albiceps attacks C. megacephala larvae during dispersal and keeps an aggregated pattern close to the release point, in single and double species, independently of the different photophases. Chrysomya megacephala single species exhibited the same pattern, but in double species this was changed to a random distribution.     

Effect of dimethoate on the developmental rate of forensic importance Calliphoridae flies

Forensic entomotoxicology has grown to impact judicial systems in developed countries. Where the use of insects and maggots as samples in death investigations as an alternative technique, especially following degradation or loss of the conventionally used samples. Carrion flies feed on dead bodies and may ingest toxic substances found in the dead body, especially when the body was poisoned before death. The knowledge of how the chemicals interact with the insect following ingestion is crucial to forensic entomotoxicologists. The study investigated the impact of dimethoate on the life cycle of four species of Calliphoridae flies, namely Chrysomya megacephala, Chrysomya saffranea, Chrysomya rufifacies and Chrysomya indiana. Various concentrations of dimethoate (1 ppm, 2 ppm, 3 ppm and 4 ppm) were utilized in the study. The rate of development of the carrion flies showed a negative correlation with the concentration of the chemical. This paper glares at the impact of the chemicals may pose to the insects, and how analysis of such impacts can guide forensic investigations of poisoning and help the investigators to solve the crime puzzle.     

Thermal ecophysiology of seven carrion-feeding blowflies in Southern Africa

A variety of temperature thresholds for larvae, pupae, and adults of seven African species of carrion‐feeding blowflies (Diptera: Calliphoridae) was measured and compared to understand their basic thermal biology and the influence of temperature on their behaviour. Calliphora croceipalpis (Jaennicke) had consistently lower temperature thresholds than all other species tested for all larval (42.9 °C), pupal (16.6 °C), and adult (45.6 °C) stages. Larvae (50.1 °C) and adults (53.4 °C) of Chrysomya marginalis (Robineau‐Desvoidy) had higher upper lethal temperature thresholds than all other species and weighed more than all other species. Pupae and adults of both Chrysomya albiceps (Wiedemann) and Lucilia sericata (Meigen) had similar temperature thresholds, whereas Chrysomya putoria (Wiedemann), Chrysomya chloropyga (Wiedemann), and Chrysomya megacephala (Fabricius) had inconsistent rank temperature thresholds between the larval, pupal, and adult stages. With a few minor exceptions, the nervous activity, muscle activity, and death thresholds in female adult flies responded at higher temperatures than conspecific male flies for all species tested. Similarly, female adult flies weighed consistently more than conspecific male flies for all species tested, except Ca. croceipalpis. These data suggest that there is a phylogenetic component to the thermal biology of blowflies, because Ca. croceipalpis belongs to a primarily Holarctic genus and shows adaptation to that climate even though it inhabits Africa. Comparisons between these temperature thresholds and the distributions of blowfly species present on three rhinoceros carcasses suggest that blowfly larvae with high upper lethal temperature thresholds (particularly C. marginalis) dominate in interspecific competition on the carcass by raising the temperature of the amassed maggots above the thresholds of other carrion‐feeding blowflies, through metabolically generated heat.     

Cannibalistic Behavior and Functional Response in Chrysomya albiceps (Diptera: Calliphoridae)

Chrysomya albiceps is a facultative predator and cannibal species during the larval stage. Very little is known about cannibalism and prey size preference, especially in blowflies. The purpose of this investigation was to determine the influence of prey size and larval density on cannibalism by third-instar larvae of C. albiceps under laboratory conditions. Our results indicate that no cannibalism occurs by third-instar larvae on first- and second-instar larvae, but third-instar larvae do eat second-instar larvae. The functional response on second-instar larvae is consistent with Holling type II. The consequences of consuming second-, compared to first- or third-, instar larvae as well as the implications of cannibalism for the population dynamics of C. albiceps are discussed.     

The toad fly Lucilia bufonivora: its evolutionary status and molecular identification

The blow fly genus Lucilia is composed largely of saprophages and facultative myasis agents, including the economically important species Lucilia cuprina (Wiedemann) (Diptera: Calliphoridae) and Lucilia sericata (Meigen). Only one species is generally recognized as an obligate agent of myiasis, Lucilia bufonivora Moniez, and this is an obligate parasite of toads. Lucilia silvarum (Meigen), a sister species, behaves mainly as a carrion breeder; however, it has also been reported as a facultative parasite of amphibians. Morphologically, these species are almost identical, and historically this has led to misidentification, taxonomic ambiguity and a paucity of studies of L. bufonivora. In this study, dipterous larvae were analysed from toad myiasis cases from the U.K., The Netherlands and Switzerland, together with adult specimens of fly species implicated in amphibian parasitism: L. bufonivora, L. silvarum and Lucilia elongata Shannon (from North America). Partial sequences of two genes, cox1 and ef1α, were amplified. Seven additional blow fly species were analysed as outgroups. Bayesian inference trees of cox1, ef1α and a combined‐gene dataset were constructed. All larvae isolated from toads were identified as L. bufonivora and no specimens of L. silvarum were implicated in amphibian myiasis. This study confirms L. silvarum and L. bufonivora as distinct sister species and provides unambiguous molecular identification of L. bufonivora.     

Foraging behaviour by an intraguild predator blowfly, <Emphasis Type="Italic">Chrysomya albiceps</Emphasis> (Diptera: Calliphoridae)

Optimal foraging theory assumes that predators use different prey types to maximize their rate of energetic gain. Studies focusing on prey preference are important sources of information to understand the foraging dynamics of Chrysomya albiceps. The purpose of this investigation is to determine the influence of larval starvation in C. albiceps on the predation rate of different prey blowfly species and instars under laboratory conditions. Our results suggest that C. albiceps prefers Cochliomyia macellaria larvae to Chrysomya megacephala under non-starvation and starvation conditions. Nevertheless, predators gained more weight consuming C. macellaria. This result suggests that C. albiceps profit more in consuming C. macellaria rather than C. megacephala. The foraging behaviour displayed by C. abiceps on their prey and the consequences for the blowfly community are also discussed.     

Complex competitive interactions in four species of Lucilia blowflies

Abstract.

• 1 The effects of intraspecific and interspecific larval competition on larval survival, adult size, adult longevity and fecundity were quantified in four species of coexisting Lucilia blowflies: illustris, silvarum, sericata and caesar.
• 2 There was a general negative effect of increasing density on larval and adult survival, size and fecundity. Additionally, complex species-specific and frequency-dependent responses were identified, which were not expected in these biologically and morphologically closely similar species.
• 3 Lucilia illustris, the numerically dominant species in the natural community, was a superior competitor to L.silvarum at intermediate densities but an inferior competitor at high density. Such nonlinear responses may be related to differences in the life histories and larval behaviour of the species (bigger eggs and more contest-type outcome of competition in L.silvarum).
• 4 We parameterized a model of interspecific competition on a subdivided resource in an attempt to reconcile the conflicting results on larval competitive abilities and the abundances of the species in the field. Using laboratory and field-estimated parameter values the model predicted coexistence of L.illustris and L.silvarum and the observed numerical dominance of the former species. The average densities of flies in the field are limited to relatively low levels, apparently preventing L.silvarum (the superior competitor at high density) from dominating and excluding L. illustris.

     

Diversity,Daily Flight Activity and Temporal Occurrence of Necrophagous Diptera Associated with Decomposing Carcasses in a Semi-Arid Environment

The harsh conditions of the Brazilian seasonally dry tropical forest known as Caatinga pose challenges to the insects specialized in the exploitation of ephemeral resources. We investigated the diversity and daily flight activity of dipterans associated with decomposing rat carcasses in a field experiment performed in the semi-arid region of Pernambuco State, Brazil. We also analyzed the temporal arrival of adult insects on the carcasses at three stages of decomposition: early, intermediate, and advanced. We collected 1173 individuals, of which Muscidae had the highest abundance (36.5%), followed by Sarcophagidae (28.1%), Calliphoridae (25.2%), and Fanniidae (10.2%). Chrysomya albiceps (Wiedemann, 1819) (Calliphoridae), Fannia pusio (Wiedemann, 1830) (Fanniidae), Atherigona orientalis (Schiner, 1868), and Musca domestica (Linnaeus, 1758) (Muscidae) were the most abundant species. The richness reached its maximum value on the second day of decomposition, with 18 species, decreasing to 8 species on the last day of decomposition (7 days). The ecological indices of diversity, dominance, and evenness varied little among the stages. There was an overlap of most species throughout the decomposition, although the overall abundance was higher at the intermediate stage for Calliphoridae, Muscidae, and Sarcophagidae. In accordance to previous studies, nocturnal flight was rare, as approximately 8% of insects were captured at night. Our results expand the knowledge on ecological and behavioral aspects of necrophagous flies under inhospitable environments, such as the dry season in the Caatinga. The dominance of the invasive species C. albiceps reinforced here illustrates its geographical expansion towards the countryside of Northeastern Brazil.     

Behavioural response of Lucilia sericata to a decaying body infested by necrophagous insects

Numerous insect species are necrophagous, with Dipterans and Coleopterans being the most abundant on a corpse. Whether the presence of necrophageous species on a corpse affects the attraction of adult blowflies to the corpse is sparsely studied. We test the hypothesis that Lucilia sericata can discriminate the odour of a noncolonized cadaver from the odour of a cadaver on which conspecific and/or heterospecific necrophagous insects are feeding. The volatile organic compounds are collected from decomposing rats under four modalities: (i) in the absence of insects; (ii) in the presence of L. sericata adults; (iii) in the presence of Dermestes frischii adults; (iv) and in the presence of both D. frischii adults and L. sericata adults. During a multiple‐choice bioassay, blowflies are exposed to the four odour samples, and are shown to prefer the odour of a corpse where conspecific larvae are present. We also expect blowflies to avoid cadavers on which predators where present, although L. sericata are not repulsed by the odour of a cadaver colonized by hide beetles. We then compare the average quantities of all 61 volatile molecules identified, finding that the presence of necrophagous insects impacts some of them. However, none of the volatile organic compounds previously reported in the literature as being attractive for L. sericata adults are impacted by the presence of necrophagous insects. The results of the present study suggest that blowfly larvae modify the volatilome of a corpse, enabling adults to discriminate a colonized from a noncolonized corpse.     

Dispersal and Burial Behavior in Larvae of <Emphasis Type="Italic">Chrysomya megacephala</Emphasis> and <Emphasis Type="Italic">Chrysomya albiceps</Emphasis> (Diptera,Calliphoridae)

Blowflies use discrete and ephemeral substrates to feed their larva. After they run out of food, the larvae begin to disperse in order to find adequate places for pupation or additional food sources, a process named post-feeding larval dispersal. Briefly state the aspects and why they are important were studied in a circular arena of 25 cm in diameter and covered with wood shavings to a height of 40 cm allowing post-feeding dispersal from the center of the arena. Larvae of both Chrysomya albiceps and C. megacephala were used in five experiments for each species. For each pupa location, determined as distance from the center, depth, and weight were evaluated. Statistical tests were done to verify the relation between weight, depth and distance for pupation and for larvae of two species shows that the media distance is significantly different for two species and for C. megacephala this distance is greater than the distance for C. albiceps. The depth too is different for each species, as the larvae of C. megacephala buries deeper than C. albiceps. With relation of weight, there is no statistic evidence that have any difference between weights for pupation for each species.     

Quickie well done: no evidence of physiological costs in the development race of Lucilia sericata necrophagous larvae

The present study focuses on trade-offs between the development rates and their life-history traits of feeding larvae. Indeed, quick growth is considered to be vital for necrophagous insects such as larvae because they are part of a rapidly changing ecosystem and at the mercy of many predators. However, excessively quick growth can have a negative effect on other life-history traits (e.g. survivorship and body size). The blowfly Lucilia sericata (Diptera: Calliphoridae) is used in the present study because this species is frequently found on carcasses in Central Europe and is a well-known experimental model in insect physiology and ecology. Individuals are reared from first instars to adults at two different constant temperatures (i.e. 15 and 28 °C) and under three different conditions: 100 Lucilia sericata (i.e. small monospecific condition), 250 L. sericata (i.e. large monospecific) or 125 L. sericata + 125 Calliphora vicina (i.e. heterospecific). The development time, pupal and larval survival rate and pupal size are determined individually under each condition. Regarding size and development time, substantial variation is observed between the different growth conditions and within a larval group under the same conditions. However, no trade-offs between development rate and size or survival are detected. These results demonstrate that, under the range of developmental conditions tested in the present study, the quick development of L. sericata larvae does not affect their size or mortality. This developmental plasticity may explain the evolutionary success of this species, which is present in several ecosystems worldwide and dominates the fresh-carrion ecosystem.     

The role of some cyclorrhaphan flies as carriers of human helminths in Malaysia

The role of some adult flies (Diptera: Cyclorrhapha) as carriers of helminth parasites of man was studied at four sites in Malaysia: a refuse dump, where no helminth-positive flies were detected, and in three peri-domestic situations where four species of flies carried up to three types of nematodes. The dominant fly species Chrysomya megacephala (Fabricius) carried eggs of the roundworm Ascaris lumbricoides L., the pinworm Trichuris trichiura (L.) and hookworm on the adult external body surface and in the gut lumen, in association with Bukit Lanjan aborigines. Chrysomya rufifacies (Macquart) and Sarcophaga spp. also had Ascaris lumbricoides and Trichurus trichiura eggs in their gut contents. Human helminths were not recovered from Lispe leucospila (Wiedemann), Lucilia cuprina (Wiedemann) or the housefly Musca domestica L. In an urban slum area of Kuala Lumpur city, filariform larvae identified as the hookworm Necator americanus (Stiles) occurred in the intestines of the face-fly Musca sorbens Wiedemann (22 larvae per 100 flies) and of Chrysomya megacephala (4.5 larvae per 100 flies). This concentration of apparently infective N. americanus in M. sorbens, a fly which often breeds in faeces and browses on human skin, could have transmission potential.     

Is aggregated oviposition by the blow flies Lucilia sericata and Phormia regina (Diptera: Calliphoridae) really pheromone‐mediated?

When female blow flies Lucilia sericata and Phormia regina (Diptera: Calliphoridae) oviposit in aggregations on carrion, even‐aged larval offspring reportedly develop faster, and fewer are parasitized or preyed upon. The benefits of aggregated oviposition equally affect con‐ and heterospecific larvae sharing a resource. The benefits imply that female blow flies engage in coordinated, pheromone‐mediated oviposition behavior. Yet, repeated attempts to identify oviposition pheromones have failed invoking doubt that they exist. Simply by regurgitating and feeding on carrion, flies may produce attractive semiochemicals. If flies were to aggregate in response to feeding flies rather than ovipositing flies, then the semiochemical cue(s) may be associated with the salivary gland. Working with L. sericata and P. regina and using liver as a surrogate oviposition medium, we test the hypotheses, and present data in their support, that (i) gravid or nongravid females ovipositing and/or feeding on liver enhance its attractiveness to gravid and nongravid females; (ii) females respond to semiochemicals from feeding heterospecific females; (iii) females respond equally well to semiochemicals from feeding con‐ and heterospecific females; (iv) macerated head tissues of females applied to liver enhance its attractiveness; and (v) females in direct contact with and feeding on liver, but not when next to yet physically separated from liver, enhance attraction of flies. We conclude that oviposition site‐seeking females do not respond to an oviposition pheromone. Instead, they appear to coopt semiochemicals associated with feeding flies as resource indicators, taking chances that resources are suitable for oviposition, and that ovipositing flies are present.